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Mohamed Fawzy Ramadan

Editor

Fig (Ficus carica):

Production,
Processing,
and Properties
Fig (Ficus carica): Production, Processing,
and Properties
Mohamed Fawzy Ramadan
Editor

Fig (Ficus carica):

Production, Processing,
and Properties
Editor
Mohamed Fawzy Ramadan
Department of Clinical Nutrition
Faculty of Applied Medical Sciences
Umm Al-Qura University
Makkah, Saudi Arabia

ISBN 978-3-031-16492-7 ISBN 978-3-031-16493-4 (eBook)

https://doi.org/10.1007/978-3-031-16493-4

© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature
Switzerland AG 2023
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and
transmission or information storage and retrieval, electronic adaptation, computer software, or by similar
or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Dedicated to the soul of my father, Professor
Fawzy Ramadan Hassanien, and my
beloved family.
Preface

Fig has been mentioned in the Holy Quran, in the form of the divine oath, in the
words of God Almighty: “By the Fig and the Olive, and by Mount Sinai,” “Surat
Al-Tin: 1-2.” God Almighty swore to fig in this noble verse because of its great
importance.
Ficus carica L. illustrates a promising item of functional food and healthy prod-
ucts. There is a lack of books presenting complete information on the production,
processing, chemistry, products, and medical traits of figs and the plant parts (i.e.,
fruit, skin, leaves, roots, latex, and by-products). This book is considered necessary
since it covers all the information about fig.
This book project aims to create a multidisciplinary discussion forum on Ficus
carica with particular emphasis on its horticulture, post-harvest, marketability, phy-
tochemistry, extraction protocols, biochemistry, nutritional value, functionality,
health-promoting traits, ethnomedicinal applications, technology, and processing.
The impact of traditional and innovative processing on recovering high-added value
compounds from Ficus carica wastes is reported. Besides, the book discusses the
potential applications of Ficus carica in food, cosmetics, and pharmaceutical
products.
Intending to provide a comprehensive contribution to the scientific community
involved in clinical nutrition, food science, horticulture, phytochemistry, health, and
pharmacology, this book comprehensively reviews the aspects that led to the recent
advances in Ficus carica biochemistry, production, and functionality. The editor
hopes the handbook will be a rich source for researchers and developers in related
disciplines.
Book chapters have a diversity of developments in food science and horticultural
research. The book contains comprehensive chapters under main sections, namely
–– Ficus carica: Cultivation, Species, and Cultivars
–– Ficus carica: Chemistry, Functionality, and Health-Promoting Properties
–– Ficus carica: Technology, Processing, and Applications

vii
viii Preface

The editor sincerely thanks all contributors for their valuable contributions and
their cooperation. The Springer-Nature staff’s help and support, especially Daniel
Falatko and Sofia Valsendur, was essential for completing my task and is highly
appreciated.

Makkah, Saudi Arabia Mohamed Fawzy Ramadan

Description

Fig (Ficus carica L.) is one of the oldest cultivated and consumed fruits worldwide.
More than 800 varieties of the Ficus carica genus are cultivated in a warm climate.
Fig is a seasonal fruit that could be harvested twice annually and could be consumed
fresh, dried, and as a jam or juice. Fresh and dry figs are appreciated as food and for
their health-promoting impacts. The syrup is used as a remedy for mild constipa-
tion. Leaves are used as fodder for animals. Fig latex is used as a curdling agent in
dairy products.
The fruit is a rich source of health-enhancing phytochemicals (i.e., phenolics,
organic acids, vitamin E, and carotenoids). Phytochemicals are influenced by the
harvest time, variety, maturity, color, fruit part, and fruit processing. Amino acids,
organic acids, fatty acids, sterols, hydrocarbons, anthocyanins, aliphatic alcohols,
volatiles, and other secondary metabolites are found in fig fruit, latex, leaves, and
root. In addition, figs are an essential source of minerals (i.e., potassium, iron, and
calcium) and vitamins (i.e., riboflavin and thiamin).
Ficus carica fresh fruit, extracts, and isolated bioactive compounds exhibited a
broad spectrum of health-promoting traits. Different nutrients and bioactive com-
pounds, including free sugars, organic acids, tocopherols, phenolic components,
and fatty acids, were detected in the Ficus carica peel extracts. Ficus carica leaves,
roots, fruit, and latex are known for their biological and health traits, including anti-
fungal, anti-helminthic, acetyl cholinesterase inhibition, and anticarcinogenic activ-
ities. Fig is used in traditional medicine for various reproductive, digestive,
endocrine, and respiratory ailments. It is used in the gastrointestinal tract and uri-
nary tract infections. Furthermore, the fig treats ailments such as diabetes, anemia,
cancer, leprosy, liver diseases, skin diseases, and ulcers. Studies reported on the
applications of fig extracts as functional edible ingredients, clinical trials to confirm
the health effects of extracts, and the valorization of plant by-products. Ficus carica
has been included in occidental pharmacopeias (i.e., British Pharmacopoeia and
Spanish Pharmacopoeia) and therapeutic guides of herbal medicines.
The influences of processing on the F. carica product’s quality and the amounts
of individual phytochemicals in fruit were studied. Drying of Ficus carica has
proven to be a reliable preservation method. The main factors that affect the drying

ix
x Description

process are temperature and the length of the process. Freeze-drying and microwave-
drying for fig preserving are also essential technologies. Besides, the effect of post-
harvest and packing methods on fruits’ phytochemicals profile and biological traits
were of interest.
Ficus carica is of importance due to its widespread food, industrial, and medici-
nal applications. Although Ficus carica products are already commercially avail-
able in the international market, it is hard to find in the bookstore works on the
production, processing, chemistry, and properties of Ficus carica.
Key Features
• Explores the chemistry of Ficus carica phytochemicals and extracts
• Discusses Ficus carica active constituents and their health-enhancing traits
• Presents the applications of Ficus carica phytochemicals and extracts
• Authored by international scientists and industry experts
• Addresses the growing application areas, including horticulture, functional food,
clinical nutrition, pharmaceuticals, and cosmetics
Readership
• Clinical nutrition, food chemistry, biochemistry, pharmacology, and horticulture
researchers and students.
• Developers of nutraceuticals, novel food, and pharmaceuticals, as well as R&D
researchers in different sectors, apply fruits and medicinal plants.
Contents

1 Introduction to Fig (Ficus carica): Production, Processing,

and Properties�� 1
Mohamed Fawzy Ramadan

Part I Fig (Ficus carica): Cultivation, Species, and Cultivars

2 Figs in Morocco: Diversity Patterns, Valorization Pathways
and Value Chain Resilience �� 11
Lahcen Hssaini, Rachid Razouk, Aziz Fadlaoui,
and Karim Houmanat
3 Tree Genome and Diversity�� 39
Fig
Dunja Bandelj, Alenka Baruca Arbeiter, and Matjaž Hladnik
4
Genetic Diversity of Fig Varieties �� 77
Rim Ben Abdallah, Imed Othmani, Amel Lagha, and Sami Fattouch
5
Bud Structure and Evolution�� 109
Giuseppe Ferrara and Andrea Mazzeo
6
Phenotypic Variability of Fig (Ficus carica L.)�� 129
Ali Khadivi and Farhad Mirheidari
7 Morpho-Chemical Characteristics Useful in the Identification
of Fig (Ficus carica L.) Germplasm�� 175
Oguzhan Caliskan, Safder Bayazit, and Derya Kilic
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid
Climate Zone�� 193
Norma Micheloud, Paola Gabriel, Juan Carlos Favaro,
and Norberto Gariglio
9
Cultivars and Agriculture Practice of Fig (Ficus carica)�� 215
Walid Nosir

xi
xii Contents

10 Physiological Behaviour of Fig Tree (Ficus carica L.)

Under Different Climatic Conditions�� 247
Aroua Ammar, Imed Ben Aissa, Faten Zaouay, Mohamed Gouiaa,
and Messaoud Mars
11 Fig (Ficus carica L.) Production and Yield�� 259
Malek Ben Temessek, Wafa Talbi, Hana Chrifa, and Sami Fattouch
12 Defense Mechanism of Fig (Ficus carica) Against Biotic
Stresses: An Advanced Role Model Under Moraceae�� 283
Sudeepta Pattanayak, Siddhartha Das, and Suryakant Manik

Part II Fig (Ficus carica): Chemistry, Functionality

and Health-Promoting Properties
13 Chemistry and Nutritional Value of Fresh
and Dried Fig (Ficus carica)�� 313
Mohamed Fawzy Ramadan
14 Fig Seeds: Source of Value-Added Oil Within the Scope
of Circular Economy�� 321
Lahcen Hssaini
15 Fig (Ficus carica) Leaves: Composition and Functional Properties�� 339
Rashida Bashir, Samra Tabassum, Ayoub Rashid, Shafiqur Rehman,
and Ahmad Adnan
16 Fig (Ficus carica) Seed Oil�� 357
Emi Grace Mary Gowshika Rajendran
17 Composition and Functional Properties of Fig (Ficus carica)
Phenolics �� 369
Mustafa Kiralan, Onur Ketenoglu, Sündüz Sezer Kiralan,
and Fatih Mehmet Yilmaz
18 Phenolic Compounds of Fresh and Dried Figs: Characterization
and Health Benefits�� 395
Aicha Debib and Soumaya Menadi
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids �� 417
Leila Meziant and Mostapha Bachir-bey
20 Fig Minerals�� 467
Sarfaraz Ahmed Mahesar, Hadia Shoaib, Abdul Rauf Khaskheli,
Syed Tufail Hussain Sherazi, Abdul Hameed Kori,
and Niaz Ali Malghani
21
Bioactive Compounds of Fig (Ficus carica) �� 479
Senem Kamiloglu and Banu Akgun
Contents xiii

22 Fig Volatiles�� 513

Mustafa Kiralan, Sündüz Sezer Kiralan, and Onur Ketenoglu
23 Fig Enzymes: Characterization, Biological Roles,
and Applications�� 523
Hesham A. El Enshasy, Bassam Abomoelak, Roshanida A. Rahman,
Ong Mei Leng, Dalia Sukmawati, and Zaitul Iffa Rasid
24 Preventive Roles of Phytochemicals from Ficus carica
in Diabetes and Its Secondary Complications �� 539
Additiya Paramanya, Nimisha Patel, Dinesh Kumar,
Fatima Zahra Kamal, Belkıs Muca Yiğit, Priya Sundarrajan,
Prairna Balyan, Johra Khan, and Ahmad Ali
25 Composition and Health-Promoting Effects of Fig (Ficus carica)
Extracts �� 561
Toyosi Timilehin George, Ayodeji B. Oyenihi, Omolola R. Oyenihi,
and Anthony O. Obilana
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts �� 579
Nusrath Yasmeen, Gondrala Usha kiranmai, and Aga Syed Sameer
27
Composition and Biological Activities of Ficus carica Latex �� 597
Mostafa M. Hegazy, Reham Hassan Mekky, Wael M. Afifi,
Ahmad E. Mostafa, and Hatem S. Abbass
28 Extraction and Analysis of Polyphenolic Compounds
in Ficus carica L.�� 643
Babra Moyo and Nikita T. Tavengwa

Part III Fig (Ficus carica): Technology, Processing, and Applications

29 Fig (Ficus carica) Drying Technologies�� 665
Olfa Rebai, Oumayma Ghaffari, and Sami Fattouch
30
Chemistry and Functionality of Processed Figs�� 689
Asad Nawaz, Noman Walayat, Ali Hassan, Maryam Chaudhary,
and Ibrahim Khalifa
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute �� 703
Akram Sharifi, Elham Taghavi, and Sara Khoshnoudi-Nia
32 Fig (Ficus carica) Shelf Life�� 723
Elham Taghavi, Akram Sharifi, Navideh Anarjan,
and Mohd Nizam Lani
33
Use of Proteolytic Activity of Ficus carica in Milk Coagulation�� 745
Hasitha Priyashantha, C. S. Ranadheera, Tharindu R. L. Senadheera,
H. T. M. Hettiarachchi, Shishanthi Jayarathna,
and Janak K. Vidanarachchi
xiv Contents

34
The Potential of Fig (Ficus carica) for New Products�� 765
Sara Khoshnoudi-Nia, Akram Sharifi, and Elham Taghavi
35 Production and Processing: A Pakistan Perspective�� 785
Fig
Aijaz Hussain Soomro and Tahseen Fatima Miano
36 Wound Healing and Ficus carica (Fig)�� 801
Nahla A. Tayyib

Index�� 811
About the Editor

Mohamed Fawzy Ramadan is a Food Chemistry

Professor at the Department of Clinical Nutrition,
Faculty of Applied Medical Sciences, Umm Al-Qura
University, Makkah, Saudi Arabia. Since 2014, he is
Professor at Biochemistry Department, Faculty of
Agriculture, Zagazig University, Egypt.Prof. Ramadan
obtained his Ph.D. (Dr. rer. nat.) in Food Chemistry
from the Berlin University of Technology (Germany,
2004). He continued his postdoctoral research at ranked
universities such the University of Helsinki (Finland),
Max-Rubner Institute (Germany), Berlin University of Technology (Germany), and
the University of Maryland (USA). In 2012, he was appointed Visiting Professor
(100% teaching) in the School of Biomedicine, Far Eastern Federal University in
Vladivostok, Russian Federation.
Prof. Ramadan published more than 300 research papers, and reviews in peer-
reviewed journals. He also edited and published several books (Scopus h-index
is 44 and more than 6500 citations). In addition, he was an invited speaker at several
international conferences. Since 2003, Prof. Ramadan is a reviewer and an editor in
several highly-cited international journals such as Journal of Umm Al-Qura
University for Medical Sciences, eFood, Journal of Medicinal Food and Journal of
Advanced Research.
Prof. Ramadan received several prizes, including Abdul Hamid Shoman Prize
for Arab Researcher in Agricultural Sciences (2006), Egyptian State Prize for
Encouragement in Agricultural Sciences (2009), European Young Lipid Scientist
Award (2009), AU-TWAS Young Scientist National Awards (Egypt) in Basic
Sciences, Technology and Innovation (2012), TWAS-ARO Young Arab Scientist
(YAS) Prize in Scientific and Technological Achievement (2013), and Atta-ur-
Rahman Prize in Chemistry (2014).

xv
Chapter 1
Introduction to Fig (Ficus carica):
Production, Processing, and Properties

Mohamed Fawzy Ramadan

1 Fig (Ficus carica): Production, Processing, and Properties

The United Nations Sustainable Development Goals (SDGs) are recognized to offer
a sustainable world vision (https://sustainabledevelopment.un.org). “Good Health
and Well-Being” goal is associated with applying health-improving plants and
environmental-friendly processes in food chain sectors (Ramadan, 2021). Due to
their safety, fruits and vegetables have been utilized as pharmaceuticals and nutra-
ceuticals. Therefore, there is a great interest in fruits and vegetables as
phytoconstituents-rich sources for nutraceuticals and pharmaceuticals (McClements,
2019; Ramadan, 2020).
Fig (Ficus carica Linn) is considered one of the oldest cultivated and consumed
fruit tree worldwide. Fig has been mentioned in the Holy Quran, in the form of the
divine oath, in the words of God Almighty: “By the Fig and the Olive, and by
Mount Sinai”, “Surat Al-Tin: 1-2”. God Almighty swore to fig in this noble verse
because of their great importance.
Ficus is one of 37 genera (family Moraceae). Fig species of the highest commer-
cial value is Ficus carica (syn. Ficus kopetdagensis Pachom.), which consists of
varieties with high genetic diversity. About 800 varieties of the Ficus carica genus
are cultivated in a warm climate. Fresh or dry figs have been appreciated as food and
for their health-promoting impacts. Besides, Ficus carica leaves are utilized as fod-
der. The syrup is utilized as a remedy for mild constipation. In addition, fig latex is
used as a curdling agent in dairy products. Ficus carica is a seasonal fruit that could
be harvested twice annually and could be consumed fresh, dried, as a jam or juice

M. F. Ramadan (*)
Department of Clinical Nutrition, Faculty of Applied Medical Sciences,
Umm Al-Qura University, Makkah, Saudi Arabia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 1

M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_1
2 M. F. Ramadan

(Mawa et al., 2013; Barolo et al., 2014; Badgujar et al., 2014; Desa et al., 2019;
Teixeira et al., 2019; Arvaniti et al., 2019; Yao et al., 2021; Salehi et al., 2021).
Ficus carica L. is a symbol of longevity and included in the human diet since
ancient time. The fruit is a rich source of health-promoting phytochemicals (i.e.,
phenolics, organic acids, vitamin E, and carotenoids). Flavonoids and phenolic
acids are the main phytochemicals in fresh and dried fruits. Chlorogenic acid, gallic
acid, quercetin-3-O-rutinoside, rutin, and epicatechin are the main phenolic acids
and flavonoids in Ficus carica fruit (fresh or dried). Phytochemical levels are influ-
enced by the harvest time, variety, maturity, color, fruit part, and fruit processing.
The analysis of flavonoids and phenolic acids in fresh and dried fig varieties and the
distribution of these compounds between fruit pulp and skin were investigated. The
antioxidant traits of fig are correlated with its content of bioactive phenolics. Amino
acids, organic acids, fatty acids, sterols, hydrocarbons, anthocyanins, aliphatic alco-
hols, volatiles, and other secondary metabolites were found in fig fruit, latex, leave,
and root. Ficus carica is an essential source of minerals (i.e., potassium, iron, and
calcium) and vitamins (i.e., riboflavin and thiamin). In addition, fig fruits are
sodium-, fat- and cholesterol-free and rich in fibers (Mawa et al., 2013; Barolo
et al., 2014; Badgujar et al., 2014; Desa et al., 2019; Teixeira et al., 2019; Arvaniti
et al., 2019; Yao et al., 2021; Salehi et al., 2021).
Ficus carica fresh fruit, crude extracts, and isolated bioactive compounds exhib-
ited a broad spectrum of health-promoting traits. Ficus carica fruits, roots, leaves,
and latex are known for their biological and health traits, including antimicrobial,
anti-helminthic, acetylcholinesterase inhibition, and anticarcinogenic effects. Fig
has been used for several types of disorders worldwide. Fig is used in traditional
medicine for various reproductive, digestive, endocrine, and respiratory ailments. In
addition, it is used for urinary tract and gastrointestinal tract infections. Furthermore,
fig treats ailments such as cancer, diabetes, liver diseases, skin diseases, anemia,
leprosy, and ulcers. Studies reported on the applications of fig extracts as functional
edible ingredients, clinical trials to confirm the health effects of extracts, and the
valorization of plant byproducts. Ficus carica is included in occidental pharmaco-
peias (i.e., British Pharmacopoeia and Spanish Pharmacopoeia) and therapeutic
guides of herbal medicine. Therefore, the fig is a promising item in pharmaceutical
biology for the formulations of novel drugs and clinical applications (Mawa et al.,
2013; Barolo et al., 2014; Badgujar et al., 2014; Desa et al., 2019; Teixeira et al.,
2019; Arvaniti et al., 2019; Yao et al., 2021; Salehi et al., 2021).
The influences of processing techniques on the F. carica product’s quality, the
phytochemicals profile, and the amounts of individual phenolics in fruit were stud-
ied. Drying of Ficus carica has proven to be a reliable preservation method. The
main factors that affect the drying process are temperature and the length of the
process. Freeze-drying and microwave-drying for fig preserving are also essential
technologies. Besides, the effect of post-harvest and packing methods on fruits’
phytochemicals profile and biological traits were of interest. On the other side, the
impact of environmental factors on the phytochemicals content suggests the best
production area and the optimum conditions for processing. On the other hand, dif-
ferent nutrients and bioactive compounds, including organic acids, free sugars,
tocols, phenolic components, and fatty acids, were detected in the Ficus carica peel
1 Introduction to Fig (Ficus carica): Production, Processing, and Properties 3

hydroethanolic extract. In addition, Ficus carica peel extract displayed promising

antioxidant and antibacterial capacities (Mawa et al., 2013; Barolo et al., 2014;
Badgujar et al., 2014; Desa et al., 2019; Teixeira et al., 2019; Arvaniti et al., 2019;
Yao et al., 2021; Salehi et al., 2021).

2 Fig (Ficus carica) Market

Recent FAOSTAT (2022) statistics reported that the global harvested area of Ficus
carica is 281,522 ha, global yield is 44,932 hg/ha, and production is 1,264,943
tonnes. Figure 1.1 presents the world’s top producing countries of Ficus carica and

300000
273853.19

250000
203559.96
200000
Tonnes

150000
91490.48
100000 85255.04
75489.41
42684.63
50000 42309.89
38834.63
25400 23758.44

Fig. 1.1 Top producer countries and production quantities (tonnes) of fig. (FAOSTAT, 2022)
4 M. F. Ramadan

the production quantities (tonnes). Ficus carica tree is one of the oldest fruits culti-
vated in the Mediterranean area and presents an essential nutritional and economic
value because of its high consumption worldwide (Melgarejo et al., 2003; Crisosto
et al., 2011; Badgujar et al., 2014; Núñez-Gómez et al., 2021). Turkey, Egypt,
Greece, Algeria, Italy, and Spain are among the Mediterranean producers, from
which ca. 90% of fig yield is produced (Sadder & Ateyyeh, 2006; Melgarejo, 2017;
Núñez-Gómez et al., 2021). Europe, Spain is the top fig-producing country in
Europe, with about 56,600 tonnes in 2019, representing ca. 4% of the world’s pro-
duction and ca. 44% of European production (Núñez-Gómez et al., 2021).

3 Fig (Ficus carica) in the Scientific Literature

As a part of this work, a survey in the literature (Scopus and PubMed) has been
performed. Fig (Ficus carica L.) highly attracts international scientific research. A
survey and scientific literature search with the keyword “(Fig and Ficus carica L.)”
in the PubMed database (June 2022) revealed 288 documents belonging to fig
(Ficus carica L.) bioactivity, phyto-extracts, oils, bioactive constituents, and
applications.
A careful search on fig (Ficus carica L.) in Scopus (www.scopus.com) showed
that the number of documents is high (approx. 1500 till June 2022). Figure 1.2 pres-
ents the document counts on fig (Ficus carica L.) from 2001 to 2021. The annual
documents published in fig (Ficus carica L.) significantly increased from 13 docu-
ments in 2001 to 187 documents s in 2021. These numbers reflect the current

200 187
Documents by year
180

160

140 121

120 104
135
100
81
72
80 68
54 80
57
60 66
49 67
40 38 32 48
29 7 13
20 21
20

0
2021 2020 2019 2018 2017 2016 2015 2014 2013 2012 2011 2010 2009 2008 2007 2006 2005 2004 2003 2002 2001

Fig. 1.2 Scholarly output on Ficus carica from 2001 to 2021. (www.scopus.com)
1 Introduction to Fig (Ficus carica): Production, Processing, and Properties 5

Note, 12 Short Survey, 1

Conference Review, 2
Book Chapter, 12
Leer, 2
Review, 34 Editorial, 2

Conference Paper,
215

Arcle, 1249

Document type

Fig. 1.3 Distribution by types of the document on Ficus carica. (www.scopus.com)

interest in fig (Ficus carica L.) as a topic in the scientific community. Figure 1.3
presents the distribution of the types of documents in fig (Ficus carica L.), which
includes research articles (1249), conference papers (215), reviews (34), and book
chapters (12). The contributions related to the subject fields (Fig. 1.4) of Agricultural
and Biological Science (47%), Biochemistry, Genetics, and Molecular Biology
(12%), Medicine (6%), Pharmacology, Toxicology, and Pharmaceutics (6%),
Environmental Science (5%), and Chemistry (4%). Scientists from Turkey, Italy,
Brazil, USA, Iran, Spain, Tunisia, China, India, Japan, France, Egypt, and Saudi
Arabia emerged as principal authors (Fig. 1.5). Acta Horticulturae, Scientia
Horticulturae, Revista Brasileira De Fruticultura, Journal Of The Japanese Society
For Horticultural Science, Plant Disease, Journal Of Plant Pathology, International
Journal Of Fruit Science, Hortscience, Food Chemistry, Frontiers In Plant Science,
Fruits and South African Journal of Botany are the leading journals that published
scientific research on the fig (Ficus carica L.).
Economics, Econometrics and Computer Science, 7, 0% Arts and Humanies, 16, 1% Health Professions, 9, 0%
Finance, 6, 0%

Muldisciplinary, 28, 1% Neuroscience, 2, 0%

Energy, 13, 1%
Mathemacs, 6, 0%
Social Sciences, 29, 1%
Business, Management and
Accounng, 3, 0%
Physics and Astronomy, 14, 1%

Decision Sciences, 1, 0%
Veterinary, 22, 1%

Earth and Planetary Sciences,

30, 1%
Nursing, 24, 1%

Immunology and Microbiology,

99, 4%

Materials Science, 28, 1%

Engineering, 48, 2%
Chemical Engineering, 58, 2%
Agricultural and Biological
Sciences, 1115, 47%
Pharmacology, Toxicology and
Pharmaceucs, 132, 6%

Medicine, 152, 6%

Chemistry, 105, 4%

Environmental Science, 124, 5%

Biochemistry, Genecs and

Documents by subject area
Molecular Biology, 293, 12%

Fig. 1.4 Distribution by subject area of documents on Ficus carica. (www.scopus.com)

Documents by country
Serbia 10
Jordan 11
Slovenia 17
Russian… 17
Indonesia 17
Germany 17
South Korea 18
Iraq 18
Portugal 23
Mexico 23
Greece 27
Croaa 28
Algeria 31
United… 34
Morocco 35
Pakistan 38
Malaysia 39
Saudi Arabia 48
Israel 49
Egypt 60
France 62
Japan 72
India 87
China 92
Tunisia 99
Spain 100
Iran 100
USA 101
Brazil 112
Italy 145
Turkey 177
0 20 40 60 80 100 120 140 160 180 200

Fig. 1.5 Distribution by country of documents on Ficus carica. (www.scopus.com)

1 Introduction to Fig (Ficus carica): Production, Processing, and Properties 7

4 Aims and Features of This Book

Fig (Ficus carica L.) illustrates an excellent functional food and healthy product
item. To the best of knowledge, there is a lack of books discussing complete infor-
mation on the production, processing, chemistry, products, and medical properties
of figs and the plant parts (i.e., fruit, skin, leaves, roots, latex, and byproducts).
Therefore, this book is considered necessary since it will cover all the information
about fig (Ficus carica L.). In addition, to the best of knowledge, this book presents
and collects available scientific information mentioned above in one work.
Fig (Ficus carica): Production, Processing, and Properties create a multidisci-
plinary discussion forum on Ficus carica with particular emphasis on its horticul-
ture, post-harvest, marketability, phytochemistry, extraction protocols, biochemistry,
functionality, nutritional value, health-promoting traits, ethnomedicinal applica-
tions, technology, and processing. The impact of processing (traditional and innova-
tive) on recovering value-added compounds from Ficus carica byproducts is
reported. Also, the book discusses the novel applications of Ficus carica in food-
stuffs, cosmetics, and pharmaceutical products.
The tentative manuscripts have a diversity of developments in nutrition, food
science, and horticulture research. The book contains comprehensive chapters under
main sections, namely
Part I: Ficus carica: Cultivation, Species, and Cultivars
Part II: Ficus carica: Chemistry, Functionality, and Health-Promoting
Properties
Part III: Ficus carica: Technology, Processing, and Applications

References

Lahcen Hssaini, Rachid Razouk, Aziz Fadlaoui, and Karim Houmanat

Abbreviations

ABTS Ethylbenzothiazoline-6-sulfonic acid

CE Catechin equivalent
cy-3 rutinoside Cyanindin-3- rutinoside
DPPH 2,2-diphenyl1-picrylhydrazyl
dw dry weight
FRSA Free radical scavenging activity
GAE Gallic acid equivalent
IC50 Half maximum inhibitory concentration
ISSR Inter-simple sequence repeat
mM Millimole
PCA Principal components (PC) analysis
RAPD Random amplified polymorphic DNA
RFLP Restriction fragment length polymorphism
SSC Soluble sugars content
SSR Simple sequence repeat
TA Titratable acidity
TAC Total anthocyanins
TFC Total flavonoids content
TPAC Total pro-anthocyanidins
TPC Total phenolic content
TSS otal soluble solids
β-Car β-carotene

L. Hssaini (*) · R. Razouk · A. Fadlaoui · K. Houmanat

National Institute of Agricultural Research, Avenue Ennasr, BP 415 Rabat Principale,
Rabat, Morocco
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 11

M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_2
12 L. Hssaini et al.

1 Introduction

Morocco hosts large biodiversity and considerable within-species genetic diversity

(Achtak et al., 2010). Located in the northwest of Africa, between the Strait of
Gibraltar and the Mediterranean Sea to the north, the Sahara Desert to the south,
with a long coastline in the west overlooking the Atlantic Ocean and crossed diago-
nally by the Atlas Mountains moving eastwards from the Atlantic, Morocco shelters
an exciting diversity of both climates and agroecosystems hosting the second most
diverse flora in the Mediterranean basin, after Turkey. The country domesticates
around 4200 species, of which over 20% are endemic (Rankou et al., 2013). The fig
(Ficus carica L., 2n = 26) tree is undoubtedly one of the most emblematic species
of the Mediterranean landscape (Khadari et al., 2008; Hssaini et al., 2020b). Being
the earliest domesticated fruit species of the Neolithic Revolution (7000–8000 B.P),
the fig tree belongs to the Moraceae family, which is one of the most populous of all
plant genera with over 800 species (Schmitzer et al., 2011; Pourghayoumi et al.,
2017; Boudchicha et al., 2018). Ficus carica L. is thought to be native to the Middle
East and has been cultivated for millennia in close association with grapevine and
olive (Barolo et al., 2014; Pérez-Sánchez et al., 2016; Vitelli et al., 2017). In
Morocco, the fig cultivation is ancestral and is mainly planted in mountains and
family gardens (Hssaini et al., 2019b). The fig landscape is characterized by four
cultivation systems based on applied agricultural practices and varietal associations.
The first system, described as extensive, concerns plantations where no cultivation
operations other than picking are practiced. These are generally small-sized farms
located on steep slopes. The second system is semi-extensive, characterized by
maintaining several agricultural practices, including tillage, pruning, impluviums,
and caprification. These practices are generally applied for both fig trees and the
intercropping system. The third system, semi-intensive, is represented by planta-
tions that benefit, in addition to the cultivation practices adopted in the semi-
extensive system, from manual weeding, maintenance pruning, and/or contribution
of mineral or organic manure. These last two systems are the most common and are
also distinguished from the extensive system by the very high number of variet-
ies used.
The country ranks the third-largest producer of figs globally after Turkey and
Egypt, with an annual production of more than 153,472 tons, representing about
11% of the worldwide production (FAOSTAT, 2019). This production is concen-
trated mainly in the Rif region (Northern Morocco), the main fig growing area
(Hssaini et al., 2019b). Traditionally, local fig resources are defined based on qualita-
tive criteria or the proper names associated with pruning, pollination, and protection
practices that make them similar regardless of being genetically far from each other.
In the Moroccan diet, figs are consumed either fresh or dried during the whole
year but abundantly during religious occasions (Hssaini et al., 2019b). This growing
interest in fig consumption is related to the fruit’s nutritional values, as they consti-
tute an essential source of fiber, minerals (potassium, magnesium, calcium, iron,
etc.), vitamins (vitamin E, vitamin C, thiamine, and riboflavin), carotenoids
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 13

(lycopene as the most abundant, zeaxanthin, α-carotene, lutein, cryptoxanthin, and

β-carotene) and antioxidant polyphenols which are more abundant in a fig than red
wine and tea and concentrated mainly in the peel (chlorogenic acid, gallic acid, and
syringic acid). In addition, figs are also rich in flavonoids (anthocyanins, catechin,
and epicatechin), sugars (glucose and fructose, with minor traces of sucrose),
organic acids, amino acids (aspartic acid and glutamine), and volatile molecules
(Solomon & Golubowicz, 2006; Oliveira et al., 2009; Veberic & Mikulic-Petkovsek,
2015; Hernández, 2016; Hssaini et al., 2019a). Those compounds are responsible
for the fig’s pleasant taste, color, astringent flavors, and aroma (Veberic et al., 2008;
Muji et al., 2012). They are also associated with preventing coronary diseases due
to their numerous biological effects, such as scavenging unstable free radicals from
anticarcinogenic and anti-inflammatory activities (Makris et al., 2007; Çalişkan &
Polat, 2012; Palmeira et al., 2019). The aforementioned bioactive compounds’
amounts strongly depend on the genetic, geographic, and growing factors (Solana
et al., 2018; Hssaini et al., 2020b; Mirheidari et al., 2020). Figs are classified as
highly perishable commodities because their high moisture content reaches up to
80% (Saki et al., 2019; Hssaini et al., 2020c). Therefore, maintaining their freshness
has always been the adequate approach to keeping their health-promoting attributes.
However, this decision implies sophisticated storage techniques and chain distribu-
tion with considerable energy consumption.
Nevertheless, figs cannot be stored for an extended period without a significant
loss of nutritional value (Mat Desa et al., 2019). Therefore, processing remains the
best way to extend a fig’s shelf-life and preserve the nutritional quality of the end
products. Traditionally, salting, dehydration, and fermentation, however, new pro-
cessing techniques are acquiring popularity worldwide since they offer key advan-
tages such as shelf-life extension, nutritional and sensory quality preservation, and
food safety (Martins et al., 2019). In Morocco, the fig processing is an increasing
challenge because of the low interest given to this sector and the low investment in
existing infrastructure, which significantly impacts the fruits and their end products’
marketability. So far, the fig processing is limited to drying, juice, and jam, even
though the fruit has many other potentials to be exploited with a high value-added
such as seeds (Hssaini et al., 2020a). The latter has been the less valorized part of
the fruit, although some recent studies reported a satisfactory oil yield with high
unsaturation level and antioxidant potency (Icyer et al., 2016; Hssaini et al., 2020a,
b). On the other hand, improving the fig value chain resilience to changing environ-
ments and different chocks is a challenging endeavor nowadays since the sector is
poorly organized and requires a holistic approach and coordinated policies to evade
undesired chocks that may occur during specific supply chain steps or in other
related sections.
This chapter focuses on the fig diversity in Morocco, highlights its valorization
pathways, and examines the fig value chain vulnerability to shocks and the explora-
tion of solutions aiming at improving the system’s ability to adapt and to be resilient
when single or multiple shocks by mapping the fig value chain weaknesses
14 L. Hssaini et al.

alongside potential consequences of various chocks that may occur taking into
account the opportunities that can be captured during the fig value chain examination.

2 Genetic Diversity Patterns

2.1 Overview of Available Fig Germplasm

The fig tree is one of Morocco’s most diversified fruit species, with over 130 mor-
photypes, mostly seedlings clones propagated by cuttings and exotic varieties intro-
duced from neighboring Mediterranean countries (Achtak et al., 2010). The
Moroccan fig genetic resources also include various unknown types and spontane-
ous forms in coexistence with cultivated types, which are few subjected to intensive
breeding programs and therefore continue to exhibit a large genetic diversity
(Hmimsa et al., 2012). This outstanding genetic diversity results from a series of
domestication events, gene flow between wild and cultivated compartments, effects
of natural adaptive selection, human selection, and large diffusion dynamics over
long periods (Khadari et al., 2005a). The impact of these processes on diversity
depends not only on the species’ biology but also on social context, human prac-
tices, and local dietary preferences (Ater et al., 2008; Perez-Jimenez et al., 2010).
Since 2008, the local fig diversity has been enriched by some exotic varieties, which
were confirmed to be suitable for drying, particularly ‘Sarilope’ from Turkey,
‘Kadota’ from Italy, and ‘Col de Dame Blanche’ from France. These varieties were
efficient under local conditions through behavioral studies undertaken by the
National Agricultural Research Institute (INRA) in 1995 (Oukabli et al., 2003a).
Two types of edible figs are growing in Morocco: biferous (breba and main crop)
and uniferous (main crop). The local biferous types are mainly represented by
eleven cultivars, ‘Ghouddane’, ‘Ournaksi’, ‘El Khal’, ‘Ember El Khal’, ‘Fassi’,
‘Messari’, ‘Filalia’, ‘Jeblia’, ‘Hamra’, ‘Ounq El Hmam’ and ‘Beida’. The uniferous
types mainly concern ‘Nabout’, ‘El Quoti Labied’, ‘Embar Labied’, ‘Hafer El
Brhel’, ‘Chaari’, ‘Ferquouch Jmel’, and ‘Ferzaoui’ (El Hajjam et al., 2018).
Denominations of the aforementioned cultivars and others refer mainly to leaf shape
and fruit morphometric traits (color, shape, or taste) and geographic origin or the
production locality (Table 2.1). In addition, the exchange of plant material, which
was accompanied by the flows of human populations, caused varietal confusion.
This problem has led to the attribution of the same denomination to various geno-
types, although they have different pomological characteristics (homonymy), or
rather naming differently some genetically identical individuals (synonymy). The
most remarkable case is that of ‘Ghouddane’ cultivar, known as polyclones (Achtak
et al., 2009). The cases of polyclonality have also been revealed in the cultivars
‘Nabout’, ‘Chaari’, ‘Ournaksi’, ‘Hamra’, and ‘Bioudi’ (Khadari & Oukabli, 2005).
This list of polyclonal cultivars is not exhaustive due to the lack of intra-varietal
molecular authentication studies within other local fig ecotypes. Due to the
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 15

Table 2.1 Denomination meaning of some fig cultivars in Morocco

Cultivar Denominations meaning
Nabouta Generic Arabic word meaning a seedling plant
Gouddaneb Dark skin color
Zerquib Blue skin color
Sebtib Originating from Sebta in northern Morocco
Messarib Originating from Beni Messara in northern Morocco
Jaadib fruit with ribs over the skin
Ounq El Refers to the pigeon neck shape
Hmamb
Ferquouch Refers to the camel foot in Moroccan dialect, which describes a typical fruit
Jmelb shape
Hmimsa et al. (2017)
a

Hssaini et al. (2019b)

environmental effect, this confusion in the cultivar denomination is linked to the

fruit phenotypic variation. In particular, the human factor’s modest technicality dur-
ing the exchange and reproduction of the plant material may also explain the
observed varietal confusions (El Khaloui, 2010; Achtak et al., 2010). The morpho-
logical similarities and the inter-connections characterizing these species have
influenced the human selection pressure exerted on the individuals, mainly resulting
from seedlings, distinguished by the local term ‘Nabout’ (Hmimsa et al., 2017). The
problems of homonymy and synonymy constitute a severe challenge in the manage-
ment, use, study, and conservation of genetic resources of fig. In addition to
Morocco, this problem has been reported in several other Mediterranean countries
such as Tunisia (Chatti et al., 2003; Saddoud et al., 2011), France (Khadari, 2005b),
Spain (Sanchez et al., 2016), Turkey (Caliskan & Polat, 2012) and Italy (Ciarmiello
et al., 2015).
The Moroccan agroecosystems also include an important diversity within the
caprifigs (male fig trees), which remains less documented. This diversity is domi-
nated by spontaneous forms resulting from seedlings growing alongside water
sources and frequently encountered near the riversides, streams, irrigation canals,
and wells (Oukabli et al., 2003b; Hmimsa et al., 2012). The named types, multiplied
by cuttings, remain less diversified. Hmimsa et al. (2017) counted less than 10 well-
known denominations of caprifig in Rif areas in northern Morocco, of which the
most popular are ‘Ḥlu’, ‘Marr’, ‘Lwizi’, ‘Ahurri’, ‘Aharchiw’, ‘Ahfriw’ and
‘Azundri’. The later remains thus far less domesticated and still growing naturally
in the wild, even though few can be found near houses and family gardens. Farmers
give little importance to caprifig cultivation, although they know its role as a pol-
len source.
Morocco set up the first national ex-situ collection in 1995 at the National
Institute for Agricultural Research (INRA) experimental station in north-central
Morocco to conserve this genetic diversity while avoiding varietal confusion. This
collection comprises 160 local clones prospected with 60 exotic varieties over the
kingdom. Lately, it has been enriched by around sixty hybrids within a program of
16 L. Hssaini et al.

intraparietal diversification and around thirty caprifig trees. Thus far, a large part of
this important fig germplasm has been screened for its morphometric, physio-
biochemical, and molecular traits, of which the superior results are briefly reported
throughout the following sections of this chapter.

2.2 Fruit Morphometric Diversity

The international guide for fig tree description includes 192 morphometric traits for
the female type, among which 40 are specific for fresh fruit, while 14 are considered
highly discriminating descriptors. The latter concerns fruit shape (width/length
index and fruit shape according to the location of the maximum width), apex shape,
fruit weight, skin ground color, internal pulp color, ease of peeling, skin cracks,
ostiole width, resistance to ostiole-end cracks, fruit flesh thickness, pulp juiciness,
fruit cavity and total soluble solids (IPGRI and CIHEAM, 2003). However, the use
of morphometric descriptors to characterize the fig diversity faces two significant
problems, firstly, the subjectivity in qualitative traits description (Khadari et al.,
1995), and on the other hand, the influence of environmental conditions that induce
fluctuations in the expressions of traits as well as uncertainties in identifying syn-
onymies and homonymies, which are widespread within the local fig populations
(Giraldo et al., 2010). Despite these inaccuracies, morphometric characterization
continues to be widely used as a practical approach in fig germplasm screening in
several Mediterranean countries (Essid et al., 2017; Khadivi et al., 2018). According
to these studies, the fruit geometry, skin color, and total soluble solids were the most
discriminant morphometric traits for fig screening. Such studies were also carried
out in Morocco, in-situ, and ex-situ, which concerned the local fig germplasm.
Regarding studies carried out in-situ, the available literature illustrates that several
surveys were performed within local fig populations in some main producing areas
to highlight the extent of existing diversity. However, it is essential to emphasize
that these studies remain questionable as the environment and agricultural manage-
ment practices are highly variable and significantly affect the phenotypic expression.
In the most recent screening studies carried out by Hssaini et al. (2020a, b, c, d)
on the fig ex-situ collection previously mentioned, the large variability revealed was
attributed to the genetic factor as the edaphoclimatic conditions alongside the
orchard management practices were the same for all genotypes. This made it pos-
sible to fix the ‘genotype x environment’ interaction impact on the expression of the
morphometric and physico-biochemical trait and thus compare the genetic potenti-
alities among 135 fig accessions (Table 2.2). It is noteworthy to mention that the
abovementioned collection was planted in 2003 following a completely ran-
dom design.
The descriptive analysis highlighted a wide variability among genotypes for the
observed traits, except for the drop at the ostiole, lenticels color, seed size, and osti-
ole width (Tables 2.3 and 2.4). Visually, 9 skin colors were distinguished within the
collection, where yellow-green and light-green were the dominant ones (Fig. 2.1).
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 17

Table 2.2 List of local clones and exotic cultivars of fig in ex-situ collection screened for fruit
morphometric and biochemical traits
Local clones Exotic cultivars
1. Ahra 2870 48. Arguil_PS8 95. Abgaiti 2111
2. Aicha Moussa 2208 49. Chaari_PS15 96. Abiarous 3015
3. Amellal 50. ELQuoti Lbied_PS11 97. Bellone
4. Assel 2890 51. ELQuoti Lbied_PS20 98. Bougie
5. Ben_T1 52. ElQuoti Lbied_PS3 99. Breval Blanca 2736
6. Ben_T2 53. ElQuoti Lbied_PS6 100. Brown Turkey
7. Bioudi 2218 54. Ghani_PS2 101. Burjasot Blanca 3037
8. Bioudi 2878 55. Ghoudan_PS1 102. Cuello Dama Blanco 2233
9. Bioudie 2255 56. Ghoudan_PS17 103. Cuello Dama Blanco
10. Bousbati 2880 57. Ghoudan_PS4 104. Colle de dame blanc
11. Btitarbi 58. Jaadi_PS16 105. Conadria (Porquert)
12. Chaari 2881 59. Lamtal_PS9 106. Conidria
13. Chaari 2881 60. Mssari_PS13 107. Diamna
14. Chabaa Ourgoud 61. Nabout_PS12 108. Dottato Perguerolles
15. El Ghani 62. Nabout_PS7 109. Figue de Marseille
16. EL Hmiri 2224 63. Ounq Hmam_PS14 110. Grise St. Jean
17. EL Khal 2283 64. Sebti_PS10 111. Grosse dama Blanca
18. EL Qoti Lezreq 2883 65. Tabli _PS19 112. Grosse Dame Blanche 2953
19. EMBAR EL KHAL 2247 66. Tabli_PS18 113. Gulgium
20. EMBAR LEBIED 2240 67. Zerqui_PS5 114. Herida
21. Fassi 68. INRA 1301 115. III 31 Roger
22. Fassi 2267 69. INRA 1302 116. Khelema 3148
23. Filalia 2211 70. INRA 1303 117. Kodata
24. Hafer El Brhel 71. INRA 1304 118. MELISSOSYKI 3074
25. Hafer Jmel 2253 72. INRA 1305 119. Nardine
26. Hamra 73. INRA 1306 120. Palmeras
27. Hamra 2252 74. INRA 1308 121. Pingo de Mel
28. Hamra 2588 75. INRA 1314 122. Princesse
29. Hmidi 2250 76. INRA 1502 123. Rey Blanche
30. Kahoulta 2251 77. INRA 1503 124. Royal Blanck
31. Lamandar Noir 78. INRA 1506 125. Snowden
32. Lmandar Bied 79. INRA 1606 126. Sucre vert
33. Mendar 2891 80. INRA 2101 127. Sucre vert
34. Nabout 2893 81. INRA 2103 128. Tena
35. Noukali 2254 82. INRA 2105 129. Trojana
36. Noukalli 83. INRA 2201 130. VCR 153/17
37. Rhoudane 2227 84. INRA 2201 131. VII 1 Roger 3
38. Taranimt 2399 85. INRA 2204 132. Violette d’Agenteruil
39. V12 86. INRA 2205 133. White Adriatic_102
40. V2 (b) 87. INRA 2206 134. White Adriatic_13
41. V33(b) 88. INRA 2304 135. Palmares
(continued)
18 L. Hssaini et al.

Table 2.2 (continued)

Local clones Exotic cultivars
42. Zerouali 89. INRA 2305
43. Ztaniti 90. INRA 2307
44. INRA 2506 91. INRA 2307
45. INRA 2508 92. INRA 2401
46. INRA 2603 93. INRA 2404
47. INRA 2802 94. INRA 2501

Table 2.3 Descriptive analysis of qualitative morphometric traits within the ex-situ collection of
INRA (135 fig cultivars)
Descriptor Dominant character Frequency ANOVA signification
Fruit shape Globose 76% **
Fruit symmetry Ovoid 70% **
Shape around stalk Rounded 55% **
Stalk abscission Medium 60% **
Ostiole color White 51% **
Drop at the ostiole Absent 65% ns
Color of ostiole liquid Pinkish 61% **
Skin color Green-yellow 46% **
Skin ribs Intermediate 81% **
Easy of peeling Medium 51% **
Skin cracks Cracked skin 72% **
Juiciness Little juicy 74% **
Skin over color Yellow 54% **
Shape of over color Absent 69% **
Lenticels quantity Scarce 43% **
Lenticels color White 66% ns
Lenticels size Medium 80% *
Pulp color Pink 45% **
Pulp texture Medium 52% **
Fruit cavity Small 52% *
Seed quantity Intermediate 50% **
Seeds size Medium 78% ns
* Significant at 0.05; ** Significant at 0.01; ns non-significant

However, according to the International Commission on Illumination (CIE), the

color coordinates of fruit skin showed much wider variability than the visual assess-
ment. They made it possible to detect color gradients varying from bright yellow to
black by revealing various intermediate colors (yellow-green, brown, purple, blue-
purple, etc.). Most studied genotypes have a globose and symmetric shape of fruit
and did not have a drop at the ostiole, with a considerable variation in fruit weight,
from 12.40 to 87.03 g. The highest fruit weights were recorded in the local geno-
types ‘Lmandar Noir’ (87.03 g), ‘Bougie’ (76.9 g), and ‘Chaaba Ourgoud’ (73.7 g)
as well as in the imported varieties ‘Kadota’ (61.5 g) and ‘Brown Turkey’ (60.4 g).
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 19

Table 2.4 Descriptive analysis of quantitative morphometric and biochemical traits within the
ex-situ fig collection of INRA (n = 135 cultivars)
Std. ANOVA
Range Mean deviation signification
Fruit weight (g) 12.40–87.03 36.31 4.784 ***
Fruit length (mm) 21.34–48.83 34.41 5.89 ***
Fruit width (mm) 25.93–61.58 39.85 7.23 ***
Stalk length (mm) 2.95–16.30 8.10 4.13 ***
Stalk width 2.92–7.96 4.86 1.23 ***
Neck length (mm) 1.64–9.52 7.01 2.16 ***
Neck width (mm) 2.22–15.69 8.64 2.62 ***
Ostiole width (mm) 2.34–8.79 5.13 1.14 ns
Fruit peel thickness (mm) 1.32–8.35 3.74 1.64 ***
Volume (mm) 10.00–85.00 35.17 14.70 ***
L* 16.71–83.64 52.36 19.32 ***
c* 1.98–63.70 27.24 15.3 ***
h* 1.24–360 120.23 25.51 ***
Soluble Sugars (g/100 gdw) 10.08–15.10 12.08 1.26 ***
Phenols (mg GAE/100 gdw) 25.33–322.00 142.74 9.78 ***
Flavonoids (mg CE/100 gdw) 14.59–103.71 42.04 9.06 ***
Anthocyanins (mg cy-3 rutinoside/100 0.41–47.95 13.57 10.3 ***
gdw)
Proanthiocyanidins (mg/100 gdw) 0.18–8.51 2.24 1.82 **
DPPḤ (mM Trolox eq/100 gdw) 20.92–488.2 180.74 10.3 ***
ABTS (mM Trolox eq/100 gdw) 31.32–658.21 425.5 14.97 ***
β-Carotene (mM Trolox eq/100 gdw) 73.5–824.31 403.42 17.38 ***
Titrable acidity (g citric acid/100 g 0.07–1.66 0.77 0.4 ***
juice)
TSS (%) 8.00–48.00 19.68 5.74 ***
Maturity index 8.79–236.72 36.08 12.52 ***
Titrable acidity (g citric acid / 100 g of 0.07–1.72 0.76 0.31 ***
juice)
pH 2.51–6.6 4.41 0.91 ***
ns non-significant; **; *** denote significant of difference at level 0.01 and 0.001respectively
GAE Gallic acid equivalent, CE Catechin equivalent, TSS Total soluble solids

However, the lowest was recorded in the genotypes “INRA 1303”, “Ounq Hmam_
PS14”, “Zerqui_PS5”, “Ghoudan_PS1”, and “Gulgium”, for which fruit weight
ranged between 12.4 and 16.4 g. The chemical screening highlighted a wide diver-
sity in the sweet taste degree of fruit, determined by the balance between total solu-
ble solids (TSS) and titratable acidity (TA). According to TSS/TA ratio (maturity
index), the sweetest figs were of the local genotypes ‘Sebti_PS10’, ‘Noukali 2254’
and ‘Chaari_PS15’ with respective ratio values of 219.30, 138.35, and 104.50,
while the most acidic were of ‘Dottato Perguerolle’ (7.85), ‘V2 (b)’ (8.81) and
‘Hafer Bghal’ (8.81). Overall, the variations of the whole observed morphometric
traits were in agreement with those reported in other similar studies carried out in
20 L. Hssaini et al.

Fig. 2.1 Fruit skin colors identified visually within the ex-situ fig collection of INRA (n = 135
cultivars). The percentage of each color within the collection is given in parentheses

other Mediterranean countries, with some differences related to the analyzed geno-
types and the environmental conditions of the experimental sites (Ateyyeh &
Sadder, 2006; Caliskan et Polat, 2008; Aljane et al., 2008; Podgornik et al., 2010;
Jimenez, 2016).
Principal components analysis (PCA) determined the most discriminating mor-
phometric descriptors among the observed traits. Thus, ten PCs explained a total
variance of 60.2%, where 27.8% was attributed to the first three PCs. The PC1
explained 12.3% of total variance and was strongly determined by 7 morphometric
traits, namely: fruit weight (r = 0.86), fruit dimensions (r = 0.69, r = 0.85 for length
and width, respectively), stalk width (r = 0.72), neck dimensions (r = 0.58, r = 0.61
for length and width, respectively) and fruit volume (r = 0.74). The PC2 contributed
9.98% of the total variance and was influenced by 4 colorimetric traits of fruit skin,
which are visual color (r = −0.66) and chromaticity coordinates, c* (r = −0.67), L*
(r = −0.65), and a* (r = 0.58). The PC3 explained 5.51% of the total variance and
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 21

was mainly linked to fruit symmetry (r = 0.60). These 12 characters were, therefore,
the most effective descriptors to discriminate the studied fig accessions. Their cor-
relations to the first three PCs are estimated to be very strong and significant, owing
to the studied collection size and the large number of the variables involved. These
findings agree with the previously reported results in edible figs (Ciarmiello et al.,
2015; Khadivi et al., 2018). A cluster analysis was performed using Ward’s method
to identify the similarities and dissimilarities among the accessions, based on all
qualitative and quantitative variables measured. The obtained clustering showed a
high morphometric polymorphism within the fig collection, distinguishing five
clusters, mostly differentiated by fruit weight, shape, and skin color (Fig. 2.2).
Given the high number of local clones included in this study, which seems to cover
a large part of the local fig genetic heritage, this investigation provides information
on morphometric diversity amplitude in the Moroccan figs, allowing a wide diver-
sity in terms of opportunities for their exploitation and valuation.

2.3 Fruit Chemodiversity

The chemo-diversity within the Moroccan fig germplasm is thus far poorly docu-
mented and still somewhat ambiguous. However, to the best of knowledge, there is
a single exhaustive ex-situ study regarding this topic performed on the same collec-
tion (Hssaini, 2020). The ultimate objective was to produce a large database on the
quality attributes of figs in Morocco, usable across various levels: varietal selection,
agroindustry, fresh consumption, quality label development, etc., and identify dis-
criminant biochemical markers for future research purposes. According to the
internationally recognized methods, a total of 11 biochemical descriptors were
involved; namely, the total contents of polyphenols (TPC), flavonoids (TFC), antho-
cyanins (TAC), soluble sugars (SSC) and proanthocyanidins (TPAC) as well as the
antioxidant activity, assessed using three assays: (i) the DPPḤ test, (ii) the ABTS
method and (iii) the β-carotene bleaching test with their respective half-maximal
inhibitory concentration (IC50) (Hssaini et al., 2019a).
The study displayed high significant (p < 0.001) differences among genotypes,
with large variation amplitudes (Table 2.4). SSC, the main factor of taste and sweet-
ness, ranged from 9.77% to 17.7%, with an average of 12.0%. Among the local
genotypes, ‘Zerqui_PS5’, ‘El Quoti Lbied_PS3’, and ‘Rhoudane 2227’ were the
sweetest, with respective SSC values of 15.1%, 14.53%, and 14%. TPC varied
between 22 and 417.5 mg GAE/100 gdw, with an average of 142.7. It was higher in
‘White Adriatic’, ‘INRA 2304’, and ‘Ghoudan_PS1’ genotypes, where the averages
were 415, 322, and 288 mg GAE/100 gdw, respectively. However, ‘INRA 1606’,
‘INRA 1503’, and ‘Rey Blanche’ were the least rich in polyphenols, with average
contents of 25.33, 28.67, and 40.89 g GAE/100 gdw. TFC was higher in ‘White
Adriatic_102’ (103.7 mg CE/100 gdw), ‘INRA 1301’ (101 mg CE/100 gdw), and
‘INRA 1306’ (96.05 mg CE/100 gdw), while lower in ‘Kadota’, ‘Rey blanche’ and
‘Diamna’, with the average TFC values were 14.6, 14.5 and 15.35 mg CE/100 gdw,
22 L. Hssaini et al.

Fig. 2.2 Cluster analysis of 135 fig genotypes based on fruit morphometric (a) and biochemical
(b) traits using Ward’s method. Discriminated clusters were marked in different colors
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 23

respectively. TAC ranged between 0.45 and 57.47 mg cy-3-rutinoside/100 gdw, with
an average of 13.57 mg. The highest values were observed in ‘INRA 2105’
(52.95 mg), ‘Burjasot Blanca’ (42.57 mg), and ‘INRA 1302’ (40.92 mg), while the
lowest were found in ‘Sucre vert’ (0.45 mg) and ‘INRA 2307’ (2.69 mg).
Nevertheless, TAC was extremely low in some cultivars such as ‘Felalia’ and
‘Trojana’ to the point that it was not detected. This was also the same case for
TPAC, which was not detected in some accessions, including ‘INRA 1314’ and
‘Princesse’. TPAC was low within the studied genotypes (0.18–10.0 mg/100 gdw).
Thus, ‘Mendar Noir’ (8.51 mg/100 gdw) and ‘Cello Dama Blanca’ (6.64 mg/100 gdw)
were the highest genotypes.
The antioxidant activity varied greatly across genotypes according to the three
used assays. Indeed, the difference between the extreme values was 23 times for the
DPPḤ assay, with an IC50 of 0.38–27.28 μg mL−1, 21 times using the ABTS test
allowing an IC50 of 0.20–5.78 μg mL−1, and 11 times according to the β-carotene
bleaching methods, for which IC50 ranged between 3.58 and 22.7 μg mL−1. Overall,
the observed variations agreed with similar previous studies (Oliveira et al., 2009;
Wojdyło et al., 2016; Sedaghat & Rehemi, 2018). Thus, some genotypes were dis-
tinguished by their high antioxidant potential, such as ‘Kahlouta’ (488.2 mM Trolox
eq/100 gdw), ‘INRA 2501’ (451.36 mM Trolox), and ‘INRA 2603’ (433.5 mM
Trolox) for the DPPḤ assay, ‘V12’ (658.2 mM Trolox) and ‘El khal’ (632 mM
Trolox) for the ABTS method and ‘Melissosyki’, ‘Hmidi’ and ‘El Qoti Lezreq’
according to the β-carotene bleaching test. The highest values of the radical scav-
enging capacity were related to the genotype richness in phenolic compounds, par-
ticularly flavonoids and anthocyanins.
According to PCA based on correlation coefficients, six traits seemed to have the
highest impact on discrimination between genotypes in terms of biochemical pro-
prieties, namely, TFC, TAC, TPAC, radical DPPḤ scavenging activity
(DPPḤ-RSA), IC50 of DPPḤ radicals (IC50-DPPH) and IC50 using the β-Carotene
bleaching test (IC50-β-Car). These traits presented correlation coefficients of more
than 0.5 to the first three PCs, with a cumulative variance of 40.91% (65.0% for six
PCs). Indeed, TAC (r = 0.73) and DPPḤ-RSA (r = 0.58) were the most correlated to
PC1, explaining 21.6% of the total variance. TFC (r = 0.81) and IC50-DPPḤ
(r = 0.70) were more linked to PC2, which contributed 10.0% of the total variance,
while TPAC (r = 0.72) and IC50-β-Car (r = 0.69) determined mostly PC3 (9.19%).
Moreover, Ward’s method’s multivariate analysis highlighted a high polymorphism
in fruit biochemical traits within the studied collection. Indeed, five clusters of gen-
otypes were obtained, mainly distinguished by soluble sugar content, free radical
scavenging activity (DPPḤ and ABTS), and content in polyphenols, particularly
anthocyanins (Fig. 2.3). Further details on phenotypic pooling and involved vari-
ables discriminating strength could be found in the studies carried out by Hssaini
et al. (2020a, b). Therefore, it emerges from this study that the analyzed fig acces-
sions, including local genotypes, exhibit a wide diversity in fruit quality, thus offer-
ing multiple choices for clonal selection and commercial and industrial purposes.
24 L. Hssaini et al.

Fig. 2.3 Ripened figs pathways beyond the farm gate. (Source: Authors’ construction)

2.4 Molecular Fingerprinting

On fig species, molecular fingerprinting investigations were carried out for a dual
objective: (i) to categorize genetic similarities between genotypes and (ii) to iden-
tify the widely encountered cases of homonymy and synonymy. In Moroccan fig
germplasm, molecular markers were fairly applied, including RFLP on mitochon-
drial DNA, RAPD, ISSR, and SSR (Khadari et al., 1995, 2005b). In the fig genetic
diversity investigation of Khadari et al. (2004), 75 accessions collected from north-
ern Morocco were analyzed using 8 ISSRs markers and 6 SSR loci. As a result, 52
genotypes, including 6 caprifig trees, were identified as genetically heterogeneous
cultivars. The homonymy cases were frequently observed in 6 denominations of
female figs, namely, ‘Bioudi’, ‘Chaari’, ‘El khal’, ‘Hamra’, ‘Ournakssi’, and
‘Rhoudane’. In addition, the SSR markers, using 17 microsatellite loci, were applied
to assess the genetic diversity within an ex-situ collection of INRA Morocco, which
includes 75 accessions representing eight caprifigs, 51 local accessions, 11 exotic
varieties, and 5 accessions of unknown origins (Achtak et al., 2009). Among the
analyzed samples, 62 distinct genotypes were identified, classified into 54 female
figs and 8 caprifigs. This study confirmed the high frequency of homonymy for the
abovementioned female figs and highlighted synonyms for 7 genotypes, namely,
‘Jeld Elhmar’, ‘Bousbati’, ‘El Har’, ‘Hamra’, ‘Fassi’, ‘Noukali’ and ‘Jeblia’.
These molecular analyzes, covering a large part of the local morphotypes, high-
lighted the high genetic diversity magnitude within the Moroccan fig germplasm
and thus confirmed the phenotypic variation revealed by multivariate analysis of
fruit morphological and biochemical traits. Some suspected cases of varietal confu-
sion were also confirmed. However, the correlations between molecular and pheno-
typic traits remain unstudied to the best of knowledge. Such studies will significantly
interest the early selection of efficient fig cultivars among local clones and created
genotypes through hybridization or mutagenesis.
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 25

3 Valorization Pathways

There has been a notable increase in fig production in Morocco during the last
decade. Thus, the production increased by about 29%, as it rose from about 109,200
tons in 2009 to 153,472 tons in 2019 (FAOSTAT, 2019). This increased rate is
mainly related to the extension of area harvested from 47,600 to 62,969 ha during
the same period, equivalent to 24.4%, as targeted by the national green Moroccan
plan launched in 2008. A large amount of this production is usually oriented to pro-
cessing, as fresh figs storage requires specific low-temperature facilities, which has
been thus far difficult to maintain throughout the distribution chain. Moreover, fresh
fig is very perishable under the ambient condition as it contains a large amount of
moisture, up to 80% (Mat Desa et al., 2019). Therefore, fresh figs can be stored for
up to 8 days under low temperature and high hygrometry (4.44–6.11 °C and 75%,
respectively). However, they can only last for 2 days under ambient storage condi-
tions (Mat Desa et al., 2019). Therefore, an extensive storage period of fresh figs
could only be achieved through continuous freezing, which requires a significant
investment and a considerable amount of energy, which is not feasible on a large
scale (Hssaini et al., 2020c).
Figure 2.3 describes the Moroccan fig valorization pathways beyond the farm
gates. Indeed, ripened figs that fail to meet the standards to direct consumption or
manufacture, mainly if they present visual blemishes, injuries, bruises, or defects,
are mainly exploited for alcoholic spirits or vinegar production or even used as ani-
mal feed. Figs meeting the aforementioned standards, including good physical
appearance and texture with hygienic conditions, are selected for direct consump-
tion or processing. For fresh consumption, several criteria are considered related
mainly to morphological characteristics such as shape, color, and caliber, alongside
taste and texture.
In Morocco, drying is the most practiced technique for fig shelf-life extending
(Chimi et al., 2008). It is also one of the apparent techniques to meet the market
demand by reducing wastage, mainly owing to the poorly established logistics for
low-temperature distribution and handling facilities leading to improper sell realiza-
tion (Mat Desa et al., 2019). Direct sun-drying along with solar-assisted drying is
particularly the main employed methods to extend fig shelf-life since Morocco
receives an intensive daily average solar radiation intensity of 5.3 kW h/m2 and
5–6 h of sunshine duration in the winter and 9–10 h during the summer (Kousksou
et al., 2015). Drying depicts a complex process of heat and mass transfer of remov-
ing moisture from biological products. In food processing, the leading role of drying
technologies is to preserve agricultural commodities’ quality, extend their shelf life
and produce new products that would not otherwise be feasible (Nurlaila et al., 2019).
Furthermore, drying allows to obtain the desired moisture content, physical
form, flavor, color, or texture and reduces the volume or the weight for low-cost
transportation (da Silva et al., 2012). In the Moroccan context, using drying is tech-
nically convenient, and the cost associated with processing, packaging, transporta-
tion, and storage is sensitively less for dried figs than for canned and frozen products
26 L. Hssaini et al.

(Rahman et al., 2016). Unfortunately, as far as we know, there are very few reports
on drying kinetics and thermodynamic studies and their impact on the physical and
biochemical attributes of figs growing in Morocco. Therefore, how different drying
techniques impact the fig drying curve and its functional properties still need to be
investigated. Also, energy efficiency and the simulation of fig drying curves with the
help of empirical and semi-empirical models are valuable for designing new or
improving existing drying systems. They are also helpful in drying process configu-
ration and control (Adak et al., 2017).
In northern Morocco, the main fig cultivation area, open sun drying remains
dominant compared to indirect solar drying techniques, as it is the most employed
by local small-holder farmers and traditional cooperatives, which dominates the
agricultural landscape. However, despite being an eco-friendly method, open sun
drying might be challenging due to the occasional occurrence of wet periods during
the fig drying period from mid-August to the last decade of September (Ekechukwu,
1999). In addition, this drying method is the slowest among other drying modes,
leading to extending the fig’s exposure to sunlight and wind. Furthermore, this
lengthy process under ambient conditions has enormous drawbacks, some of which
are related to the high probability of peel browning occurrence, microbial growth,
pest disturbance, insect infestation, and dust contamination, which are disadvanta-
geous to the final product quality. Furthermore, in previous studies, direct sun-
drying of several fruits and beverages is associated with postharvest product loss,
which is more problematic in developing countries (Mesta et al., 2013; Bradford
et al., 2020; Jayaraman & Gupta, 2020). Today, there is growing awareness of the
need to adopt an eco-friendly drying mode that fulfills market demands for eco-
nomic value-added and consumer satisfaction by producing hygienic dried figs with
high nutritional properties (Durance, 2002). Solar dryers can fulfill the abovemen-
tioned requirements and substitute direct sun-drying techniques (Chua & Chou,
2003). Unlike direct solar dryers that use solar radiation, solar dryers use heated air
in active and passive circulation modes. For figs, these dryers can reach a tempera-
ture of 50–60 °C and thus produce a good product quality under constant tempera-
ture and moisture removal rate (Singh et al., 2018).
In Morocco, the overall quality of dried figs quality is most often determined
through several key attributes related to the physical appearance (bright color and
optimal hygienic aspect), chemical (high sugars, optimal water activity, and tex-
ture), and microbial (absence of microbial load, pests, and contaminants). However,
some issues can occur during or after drying, diminishing the product quality and
marketability (Tan, 2017; Aksoy, 2017). Peel browning related to enzymatic or
Maillard reactions is among these issues which concern white peel-colored figs.
Thus, the color change is the primary limitation on dried fig’s shelf-life that influ-
ences consumer acceptance (Hssaini et al., 2020c). Enzymatic browning is caused
by the conversion of polyphenols to quinones, which is catalyzed by polyphenol
oxidases, while Maillard browning is a result of the crystallization of carbohydrates
from inside to the external surface of the fruit (Tareen et al., 2012; Gamboa-Santos
et al., 2014; Derardja et al., 2019; Mat Desa et al., 2019).
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 27

Moreover, both browning types speed up as the temperature increases (Bet,

2015). Several approaches are employed to deal with these issues, including the
application of pretreatments which are divided into three groups: chemical, using
organic acids, such as exogenous ascorbic and salicylic acids (Hssaini et al., 2021a),
physical like using heat such as blanching in boiling water (Piga et al., 2004;
Piazzolla et al., 2018) or by irradiation (i.e., gamma rays and ultraviolet) (Hamanaka
et al., 2011; Afsah-Hejri et al., 2021), or finally biological through the application
of plant extracts particularly aromatic and medicinal plants (Allegra et al., 2017,
2021). In Morocco, sulfites are well-known anti-browning agents and thus are
extensively used to prevent browning reactions, despite their negative effect on con-
sumer health (Oliphant et al., 2012; Stohs & Miller, 2014). Efforts are deployed to
substitute sulfites with more healthy and safe anti-browning chemicals such as
ascorbic and salicylic acids (Hssaini et al., 2021a). Considering those above, some
consumers prefer dark-colored fruits, as they do not need a few additional treatments.
In the scope of circular processing, unsold dried figs are used either for wine or
spirits production, with plurivarietal or monovarietal dried figs due to their reduced
quality due to storage conditions or time or even both the raw inputs (Solana et al.,
2018). They could also be roasted and used as a coffee substitute or animal feed
(Ishurd et al., 2004; Liang et al., 2016; Mat Desa et al., 2019). Processing to jam and
syrup comes second after drying but in few amounts and, similar to some other
products, are in some cases meld with extracts of medicinal and aromatic plants,
especially thyme (Allegra et al., 2017, 2021). Recently, some research outlined the
great opportunity behind fig seeds processing as a novel source of value-added oil
with a high unsaturated oil yield reaching up to 30% and presenting a promising
antioxidant potency (Hssaini et al., 2020a, 2021b). Finally, as the third worldwide
fig producer, Morocco still has millstones to achieve in fig processing as it hosts a
wide range of diversity with excellent attributes that need to be further valorized so
that the Moroccan fig and its derivates could be competitive in the worldwide market.

4 Value Chain and Its Resilience

In Morocco, no single study is interested in studying and assessing the fig value
chain, probably because the sector remains poorly structured and lacks reliable data.
Nevertheless, the fig supply chain value in Morocco is depicted in Fig. 2.4, based on
observations made in the field. The figure draws a traditional food supply chain
model, where vertical connections are spotted within the interactions between
stakeholders and system operations. At the model upstream end, the classical supply
chain is mainly driven by smallholder farmers and agroindustry local-oriented, pro-
viding fresh figs and its fig-based products. The main way fresh figs and processed
ones reach the traditional markets and supermarkets is through a web of stakehold-
ers, including small and medium-scale traders and trailers who purchase figs at the
farmgate or traditional markets and transport these commodities to either rural or
modern fresh fruits markets or local cooperatives for drying or producing traditional
28 L. Hssaini et al.

Fig. 2.4 Schematic representation of figs supply chains in Morocco, with potential outcomes and
disturbances drivers. (Source: Authors’ construction)

fig-based preparations. At the model downstream end, figs and fig-based products
reach consumers through rural assembly markets and supermarkets.
Going back to the model upstream end, small and medium-scale agro-industrial
firms provide consumers through modern markets with fig-based products. It is
noteworthy that the market share of figs and fig-based products in the supermarkets
remains, thus far, much smaller than the traditional markets. Export-oriented supply
is mainly based on medium retailers who provide export-oriented agro-industry for
fresh and dried figs. Between 2005 and 2020, these exports were 46 tons (t) for fresh
figs, 455 t for dry figs for human consumption, and 250 t for damaged dry figs (com-
pletely discarded for not meeting consumption standards) for industrial use.
Morocco imports mainly dried figs for human consumption through a tiny network
of local and foreign importers, mainly from Turkey, the first worldwide dried fig
exporter (FAOSATA, 2019). Imported figs are subjected to import duty set at 40%
upon entry into the country (Moroccan Office of Change, 2021). These commodi-
ties are also subjected to value-added tax at a rate of 20% and the parafiscal import
tax of 0.25%. However, thanks to free trade agreements, some countries are exoner-
ated from these duties.
Based on the model depicted in Fig. 2.4, many weaknesses persist upstream and
downstream, hindering its development. At the upstream level, the sector is subject
to structural constraints, as it is characterized by the dominance of small farms,
excessive fragmentation, and the multitude of legal systems in the fig landscape.
Furthermore, most orchards are managed traditionally with old-aged trees. Except
for some experiences that are beginning to emerge, the professional organization of
producers is still very weak, limiting the possibilities to access production key fac-
tors at reduced prices and technical assistance services for modernization of man-
agement methods. At the midstream level, the markets for fig products have an
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 29

oligopolistic nature, characterized by numerous small and medium-sized producers,

who are poorly organized alongside a few large buyers who set prices. Thus, more
than a third of the sector’s total value added is captured by intermediaries. Besides,
the insurance of fig producers regarding the impact of climate change risk appears
to be limited only to six hazards, namely hail, frost, strong wind, dry-hot wind,
overheating, and excess water. However, it does not cover the drought, pests, and
diseases. At the downstream level, fig consumption, mainly dried figs, is still poor
and follows a stagnant rate of about 0.15 Kg per capita per year (High-Commission
for Planning of Morocco, 2014). Notably, the dried figs consumption rate follows
the demographic growth besides being mainly concentrated during religious events,
including the fasting month of Ramadan (Achtak, 2009). The combination of all the
above mentioned factors has limited the possibilities of intensification and econo-
mies of scale, leading to the low profitability of this crop.
Some of the major limitations that face the modernization of the Moroccan fig
supply chain are the lack of organization among fig-trees, and farmers, within a
coordinated aggregative, alongside the absence of coordination between itinerant
components involved in the fig supply chain, mainly traders and retailers. Therefore,
we hypothesize that fig supply chain actors, including smallholder farmers, small
and medium-scale traders, and rural assembly markets are more vulnerable when
the fig supply chain is disrupted.
Today, there is a rising concern about bringing resilience thinking into the food
system and the fig supply chain. The global food system is increasingly complex
through transversal interdependencies across various stakeholders, locally or glob-
ally. Despite the very positive results, these diagonal interdependencies also engen-
der system vulnerabilities to many factors and conditions, together with system
shocks. According to Meuwissen et al. (2019), a food system resilience is defined as
the system’s ability to withstand disturbances and adapt to external stress over time.
According to Tendall et al. (2015), disturbances in the food system can be sudden or
gradual undesirable changes, internal or external, with structural or cyclical impact.
These disturbances can separately or jointly impact food availability, access, and
utilization. These three dimensions describe the three first components of food secu-
rity (FAO, 2008). The social and economic impact can also be considered cumula-
tive impacts of the aforementioned disturbances, where smallholder farmers and
their livelihood may be the weakest link.
Bringing resilience to the food value chain toward unexpected shocks such as the
recent crises caused by the COVID-19 pandemic, of which the impacts on human
lives and livelihoods were and continue to be devastating, is an urgent challenge.
Moroccan figs sector cannot withstand any disturbance because of the lack of strong
structure and coordination among the system components and stakeholders.
Therefore, many weaknesses can be spotted from the actual model, such as (1) lack-
ing efficient marketing strategy and insufficient development of valorization routes,
(2) small-sized farms, (3) weak coordination among stakeholders, and low connec-
tions strength in the value chain, (4) quality and quantity losses due to inadequate
post-harvest facilities and infrastructures such as cold and storage, and (5) low
30 L. Hssaini et al.

Fig. 2.5 Scheme showing the food system resilience to various shocks with their impacts over
different fig value chain levels. (Based on the model proposed by Tendall et al., 2015)

investment in new agricultural technologies. These weaknesses can broadly weaken

the fig value chain resilience and thus make it easily exposed to various shocks.
According to Simonovic and Peck (2013), the food system resilience pattern can
be depicted in four main regions, including (1) robustness, which is the capability to
withstand the disturbance occurrence for a consistent time before showing any food
security loss, and (2) redundancy, which is the capacity of absorbing disturbance
effect, adaptiveness describing the ability to recover the lost occurred in food secu-
rity (3) resourcefulness and (4) adaptability, which defines how much and how fast
the lost in food security is recovered. For the Moroccan fig value chain that lacks
tangible indicators for resilience behavior, unforeseen disturbances impacts can be
drawn following three levels: upstream, midstream, and downstream (Fig. 2.5),
according to the scheme in Fig. 2.4. Thus, for the upstream end, consequences could
be seen in production and processing disturbances besides a loss in smallholders’
livelihoods. Downstream, interconnexions between retailers and treaders can inter-
rupt figs and fig-based products transportation and storage. Disruptions can also
induce great instability in product prices and, therefore, impact social welfare. At
the consumer level, disturbances can occur in the consumption pattern and the bal-
ance between the demand and offer.
Finally, no attempt was previously made to scale the fig value chain and assess
its resilience toward unforeseen shocks, like the ongoing one engendered by the
COVID-19 pandemic. Therefore, this chapter tried to bring more attention to the
poor connections within the Moroccan fig value chain, which makes its resilience
very questionable. Bringing resilience to the fig value chain will build its ability to
bounce back when confronted with a specific disruption and allow its webs of stake-
holders and their interconnexions to return to their initial configuration. This will
safeguard the continuous flow of goods through the system and drive the supply of
fig and fig-based products. However, this is a challenging task that requires perti-
nent organization and collaborative coordination from all stakeholders.
2 Figs in Morocco: Diversity Patterns, Valorization Pathways and Value Chain… 31

5 Conclusion

Morocco hosts a fascinating fig diversity that has gone largely unheeded and is natu-
rally preserved in traditional agroecosystems, particularly in the northern regions.
Available research has mapped a large phenotypic and chemotypic diversity using
high discriminating methods. However, this diversity pattern, particularly locally
cultivated figs and wild-growing figs, know substantial homonymy and synonymy
cases. This outstanding diversity remains confusing and undocumented and deserves
to be further exploited. Even though this fruit is globally recognized as a traditional
pantry for healthy eating, Moroccan fig and its end products are less competitive on
the international market. As a highly perishable commodity, fig processing remains
less developed, contributing to low marketability. The current chapter also aims to
deepen our understanding of the resilience of fig chain value to changing environ-
ments and shocks, including the disruptions that emerged because of the ongoing
covid pandemic. This study brings significant nuances to the debate that should be
undertaken on Moroccan fig chain value resilience to various types of shocks and
explores potential solutions to the benefit of policymakers, industry, and end-users.

Acknowledgments The authors are thankful to Zahra Oussi Ali and Ali Hssaini.

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Chapter 3
Fig Tree Genome and Diversity

Dunja Bandelj, Alenka Baruca Arbeiter, and Matjaž Hladnik

Abbreviations

1-MCP 1-Methylcyclopropene
ABA Abscisic acid
AFLP Amplified Fragment Length Polymorphism
AGB Active Germplasm Bank
BERC Biodiversity and Environmental Research Center
CAPS Cleaved Amplified Polymorphic Sequence
CIHEAM International Centre for Advanced Mediterranean Agronomic Studies
CRDA Commissariat Régional au Développement Agricole
CRPh Centre de Recherches Phoénicicoles
DNA Deoxyribonucleic acid
EST Expressed Sequence Tag
EST-SSRs Short Sequence Repeats derived from Expressed Sequence Tag
FEIS Faculty of Engineering of Ilha Solteira, University of São Paulo
GA Gibberellin
GWAS Genome-wide association study
He Expected heterozygosity
Ho Observed heterozygosity
HTS High-throughput sequencing
INRA Institut National de la Recherche Agronomique
IPGRI International Plant Genetic Resources Institute
IRA Institut des Régions Arides
ISA Institut Supérieur Agronomique
ISSR Inter-Simple Sequence Repeats
ITS Internal Transcribed Spacer
KASP Kompetitive Allele Specific PCR
LTR-RE Long terminal repeat retrotransposon
mtDNA Mitochondrial DNA

D. Bandelj (*) · A. Baruca Arbeiter · M. Hladnik

Faculty of Mathematics, Natural Sciences and Information Technologies, University of
Primorska, Koper, Slovenia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 39

M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_3
40 D. Bandelj et al.

n Number of amplified alleles

NCGR National Clonal Germplasm Repository
ne Number of effective alleles
NGS Next-generation sequencing
ONT Oxford Nanopore Technologies
PacBio Pacific Biosciences
PC Principal Component
PCA Principal Component Analysis
PCR Polymerase Chain Reaction
PDO Protected Designation of Origin
PI Probability of identity
PIC Polymorphic information content
RAD-seq Restriction site-associated DNA sequencing
RAMPO Randomly Amplified Microsatellites Polymorphisms
RAPD Random Amplified Polymorphic DNA
RFLP Restriction Fragment Length Polymorphism
RNA Ribonucleic acid
SMRT Single-molecule real-time sequencing
SNP Single nucleotide polymorphism
SSR Simple sequence repeat
STMS Sequence-tagged microsatellites
TE Transposable element
TGS Third-generation sequencing

1 Origin, Taxonomy and Biological Properties

of the Common Fig

Undoubtedly, the common fig is one of the oldest fruits accompanying man and is
associated with the beginning of horticulture in the Mediterranean basin and south-
west Asia (Zohary et al., 2012). Archaeobotanic remains, dated back to
11,200–11,400 years ago, have confirmed its consumption among settlers in an
early Neolithic village in the Lower Jordan Valley (Kislev et al., 2006). Next, the fig
tree was spread from its center of origin to southern Arabia and western Asia, and
introgression of local wild figs and landraces and human selection contributed to the
development of diverse forms and varieties (Aradhya et al., 2010). Further spread-
ing of the common fig from the east to the west Mediterranean basin into Greece,
Italy, Spain, Portugal, and towards the south, into Egypt increased the number of
varieties (Aradhya et al., 2010). Genetic diversity study of mtDNA of Mediterranean
fig populations has revealed the existence of three gene pools -Balearic, West, and
East Mediterranean- and significant genetic differentiation between the West and
East Mediterranean (Khadari et al., 2005). In the mid-sixteenth century, Spanish
missionaries brought the common fig into the New World, and Franciscans were the
3 Fig Tree Genome and Diversity 41

first to plant figs in California around the mid-nineteenth century (Aradhya et al.,
2010). Finally, the fig tree was introduced from its native area to China about a
thousand years ago, where significant progress in its research has been obtained in
selecting the most suitable varieties, cultivation techniques, processing, and medical
use (Lianju et al., 2003).
The fig tree (Ficus carica L.) belongs to the mulberry family (Moraceae), which
includes about 39 genera (World Flora Online, 2021). The genus Ficus L. contains
more than 841 species, mainly grown in the tropics (World Flora Online, 2021), and
is considered one of the largest genera of flowering plants (Lansky & Paavilainen,
2011). The genus Ficus L. is currently divided into 17 sections, and the species
Ficus carica L. is placed within the section Ficus. The species F. carica is repre-
sented by two subspecies, (1) Ficus carica L. subsp. carica, known as the common
fig and native to Afghanistan, Cyprus, Greece, Iran, Iraq, Crete, Lebanon-Syria, the
North Caucasus, Palestine, Tadzhikistan, Transcaucasia, Turkey, and Turkmenistan,
and (2) Ficus carica subsp. rupestris (Hausskn. ex Boiss.) Browicz, native to
Afghanistan, Iran, Iraq, Lebanon-Syria, Pakistan, Turkey, and the Western
Himalayas. In addition to the native species, the common fig and numerous cultivars
are spread and widely cultivated in the tropics and subtropics (GRIN, 2021;
KEW, 2021).
Ficus species are gynodioecious and functionally dioecious with unique pollina-
tion biology. They have developed a mutual symbiosis with specific pollination
wasps of the Agaonidae family. The advantage of highly specific symbiosis (a single
specific agaonid wasp pollinates the majority of Ficus species) is that it allows the
development of the wasp. At the same time, the wasp’s presence allows pollination
events inside the fig syconia. F. carica has developed a symbiotic relationship with
its wasp pollinator Blastophaga psenes L., which enters the syconium through the
ostiole to oviposit short-styled female flowers. The F. carica -Blastophaga symbio-
sis system has stabilized the fruiting phenophase of the common fig, and dioecy
prevents inbreeding and is a result of adaptation to seasonal climate conditions
(Kjellberg et al., 1987, 2005; Mars et al., 2017).
The common fig possesses two tree morphs with three functional floral forms
(Fig. 3.1): (1) The edible fig produces syconia with long-styled female flowers
(female tree); (2) The caprifig produces syconia with short-styled female flowers
(serves as the oviposition site for the wasp) and male flowers (functionally male
tree) (Beck & Lord, 1988). Based on the floral biology and cropping/pollination
characteristics, figs are further classified into one of four types (Stover et al., 2007;
Flaishman et al., 2008; Çalişkan et al., 2017): (a) the common type, also called
‘persistent’, which develops fruits parthenocarpically (one or two crops), (b) the
Smyrna type, which requires pollination with pollen from (c) the caprifig (goat fig),
and (d) the San Pedro type, which produces the first crop through parthenocarpy
(breba), and a second crop (main crop) after caprification (Ikegami et al., 2013a).
The caprifig, which is also considered as wild and an ancestor form of the female fig
(Mori et al., 2017), is thus an important source of pollen in the caprification process
in which the pollen from caprifig trees is transferred to Smyrna and San Pedro types
by the pollination vector B. psenes (Çalişkan et al., 2017). Caprifigs produce three
42 D. Bandelj et al.

Fig. 3.1 Schematic representation of gynodioecious fig tree (F. carica) flower types and female fig
types differentiation based on the need for caprification to set a crop

crops during each growing season: ‘profichi’, ‘mammoni’ and ‘mamme’. Only
‘profichi’, which ripen in the early summer, are the source of pollen and the pollina-
tion occurs when female trees and caprifigs are present in the same orchard (Essid
et al., 2015).
Domestication of the fig tree has been focused on the selection and vegetative
propagation of female clones with desirable features of fruits and expressed parthe-
nocarpy. Vegetative propagation with cuttings or other vegetative parts of the plant
has preserved the original features of the maternal trees. In addition, domestication
has increased fruit size, amount of flesh, and sugar content (Zohary et al., 2012).
Fig genetic resources can be divided into cultivated, wild, and feral forms. Wild
populations of figs are privileged to a few isolated parts of native forests, such as
rock crevices, springs, wet gorges, and stream banks, and their reproduction is based
on seed dispersal. They are often complemented by feral forms, which are not
exactly wild. Feral forms derived from seeds produced by local domesticated clones
pollinated by the adjacent wild-growing caprifigs and distributed sporadically in
secondary habitats, such as the edges of plantations, ruins, collapsed cisterns, or
moist cave entrances (Zohary et al., 2012). Both feral and wild forms usually have
relatively small, hardly-edible syconia and show phenotypic plasticity with consid-
erable variation in their morphology and vegetative traits (Zohary, 1983).
Reproduction is exclusively sexual and depends on elaborate symbiosis with the
B. psenes wasp (Zohary, 1983; Galil & Neeman, 1977). After random cross-
pollination, plants developed from seeds are highly diverse with distinct character-
istics (Zohary & Spiegel-Roy, 1975).
3 Fig Tree Genome and Diversity 43

2 Fig Genetic Resources: The Importance of Research

At the end of the twentieth century, the common fig was classified as an underuti-
lized and neglected fruit species. However, in the last twenty years, its status has
changed since many published research articles are now available, and there are
many ongoing activities orientated towards the characterization and conservation of
fig genetic resources at the national and international levels.
The economic value and functional properties of common fig fruits (Vinson,
1999) and other phytochemicals with potential health-promoting effects found in
figs have contributed to its broader interest in the investigation (Arvaniti et al., 2019;
Hssaini et al., 2020a). A review of worldwide ethnomedical uses of F. carica
reported its traditional use in healing more than 40 disorders (Badgujar et al., 2014).
Phytochemicals with biological activities are found in fig leaves, fruits, latex, and
roots and places the common fig among promising candidates for the pharmaceuti-
cal industry to develop new drugs and for clinical uses (Badgujar et al., 2014).
Moreover, recent studies have shown that figs can also be used for new purposes.
The fig seed oil contains unsaturated fatty acids, gamma-tocopherol and is a rich
source of omega-3 and omega-6 fatty acids (Kucukerdonmez et al., 2021). Soltana
et al. (2015) confirmed the favorable nutritional properties and dietary importance
of oil from fig achenes. Their results have also opened up new possibilities for valo-
rizing waste material generated from fig fruit processing, which can be used in the
cosmetic industry. These findings support the integrated research of fig genetic
resources patrimony involving genomic, transcriptomic, and proteomic levels.
The genetic patrimony of the common fig has been developed through millennia
of cultivation. The unique and special pollination biology of the common fig and
early domestication and propagation practices focused on selecting genotypes based
on morphological or other prospective traits have contributed to and influenced the
development of fig genetic resources and diversity. In addition, outcrossing the cul-
tivated common fig with wild individuals and other artificial selections of interest-
ing phenotypes has contributed to common fig diversity. Conversely, the different
ecological factors, the selection criteria, and the vegetative propagation of a limited
number of fig genotypes could have led to reduced diversity and have been reported
in some fig-producing countries (Saddoud et al., 2007; Boudchicha et al., 2018).
Characterization of common fig varieties at the national and international level
contributes to understanding geographic patterns of genetic diversity distribution.
The relationships between fig varieties may be unclear due to the exchange of plant-
ing material among fig-producing countries and their export from the Mediterranean
basin to other continents. The result of these transfers is a generation of many syn-
onyms (several denominations of the same genotype) and homonyms (several geno-
types under the same denomination). Dispersal of varieties and the lax use of names
by users or just because of incorrect translation of original names (Aradhya et al.,
2010) have contributed to the variety denomination confusion. For example, the
study of fig varietal diversity and denomination in Northern Morocco has demon-
strated that out of 133 fig varieties, 39% had one or more synonyms (Hmimsa et al.,
44 D. Bandelj et al.

2012). Clear distinction of varieties is also essential for developing fig products with
quality labels such as the protected designation of origin (PDO), which provides
added value and is highly recognized by consumers. Accurate identification of vari-
eties is of essential importance during the propagation process in nurseries to ensure
the identity of the planting material and thus the production economy.
In the light of changing climate conditions, the emergence of new pests and dis-
eases, and demanding marketing standards, genetic diversity represents an essential
source for plant breeders. The common fig has not been subjected to intensive
breeding programs, and from this point of view, it is essential to fully characterize
its genetic patrimony (Perez-Jiménez et al., 2012). The main breeding programs are
oriented to achieve farm and market requests in terms of biotic and abiotic resis-
tance, the improvement of nutritional value, higher productivity, elimination of cap-
rification, the persistence of mature syconia, and bridging the gap between first and
second crop fruitfulness (Bohanec, 2008). From this perspective, phenotyping of fig
genetic resources, together with molecular characterization, is essential for the
effectiveness of breeding programs and germplasm conservation efforts.
Furthermore, millennia of cultivation, propagation practices, and environmental-
ecological factors have contributed to the development of local ecotypes in a given
geographical area. These genotypes could be the carriers or reservoirs of some
important genes related to the quality traits and resistance to abiotic and biotic stress
factors and are important to breeders looking for a unique combination of traits
or genes.
The awareness of the importance of preserving the diversity of crops contributed
to the establishment of national and international germplasm banks where all diver-
sity of the target crop is collected and maintained. The inventory of local genetic
diversity of figs in a defined geographical region is usually the first step in catalog-
ing all different ecotypes/varieties. Identified, distinct phenotypes are then clonally
propagated and planted in regional or international collections (germplasm banks),
where accessions are further characterized with morphological descriptors, agro-
nomical traits, and quality parameters under uniform ecological conditions and with
molecular markers. Gene banks or ex-situ collections, therefore, represent a unique
repository of plant diversity, which significantly contribute to effective conserva-
tion, management, and utilization of germplasm. They offer insight into the amount
and patterns of genetic diversity distribution and permit germplasm classification
based on genetic similarities and differences (Aradhya et al., 2010). The accurate
identification and valorization of collected accessions is an important task, leading
to better management of collections by identifying gaps, including deficiencies and
redundancies of the accessions. In addition, the collections contribute effectively to
the conservation of rare, local genotypes and prevent the loss of individual genes,
sets of genes, or in the extreme scenario, genetic erosion of the population of the
species.
3 Fig Tree Genome and Diversity 45

3 Tools for Fig Genetic Resources Diversity Studies

3.1 Morphological Evaluation

The traditional approach of fig germplasm characterization is based on the descrip-

tion of morphological characters and traits. One of the first major works published
in the twentieth century presenting a review and descriptions of fig varieties was the
monograph written by Condit (1955). His work was based on his observations and
studies extending over more than thirty years. In 2003, the Descriptor List for Ficus
carica L. was released by the International Plant Genetic Resources Institute
(IPGRI) and International Centre for Advanced Mediterranean Agronomic Studies
(CIHEAM) as a result of the collaborative work of scientists from different coun-
tries (IPGRI and CIHEAM, 2003). The main goal was to provide a universal, stan-
dard tool for fig genetic resources investigation. The list of descriptors is divided
into several categories. Characterization descriptors, which are usually highly heri-
table and less dependent on environmental conditions, allow easy and quick dis-
crimination between phenotypes. The list of characterization descriptors for figs
involves plant descriptors, i.e., biological characters (9 characters), growth descrip-
tors (16 characters with some subcharacters), and leaf (21 characters) and fruit
descriptors (43 characters). For evaluation of caprifig, 26 characters related to the
crop setting fruits (‘mamme’, ‘profichi’ and ‘mammoni’) were selected: fruit yield,
amount of gall flowers, presence of male flowers, pollen maturation, Blastophaga’s
exit, presence of parasitic insects, and germination rate of pollen. The list of evalu-
ation descriptors includes plant descriptors and agronomic performance, including
yield (8 characters), abiotic (7 characters), and biotic (2 characters) stress suscepti-
bility. Evaluation descriptors are more related to the information about traits needed
for crop improvement, and many of the descriptors in this category are more depen-
dent on environmental conditions. This category of descriptors also includes bio-
chemical (isoenzymes and polyphenol profiling, for example) and molecular
markers (RFLP (Restriction Fragment Length Polymorphism), AFLP (Amplified
Fragment Length Polymorphism), RAPD (Random Amplified Polymorphic DNA),
STMS (sequence-tagged microsatellites), PCR-sequencing, and others). The authors
also proposed a list of highly discriminating descriptors of figs, including fruit and
leaf characters, maturity, crop setting, need for pollination, rooting ability, yield, etc.
Although the use of DNA markers represents one of the basic approaches and
good practice in evaluating and managing plant genetic resources today, the mor-
phological description and phenotyping remain the first step in evaluating germ-
plasm, allowing the first insight into the diversity of the plant. Studies have reported
evaluation of fig genetic resources with morphological descriptors or combined
(morphological markers together with physiological, phenological and biochemical
traits) in the common fig. This information is valued in pre-breeding programs and
helps select the most appropriate genotypes with desired traits.
IPGRI and CIHEAM fig descriptors (2003), as well as some others additionally
implemented by different research groups (Giraldo et al., 2010; Podgornik et al.,
46 D. Bandelj et al.

2010), have been widely used for fig germplasm characterization in many fig pro-
ducing countries such as Turkey (Çalişkan & Polat, 2012a, b), Slovenia (Podgornik
et al., 2010), Spain (Giraldo et al., 2010), Morocco (Hssaini et al., 2020a, b),
Palestine (Ali-Shtayeh et al., 2014), Iran (Fatahi et al., 2017), and Brazil (Rodrigues
et al., 2019). Studies reported their usefulness for discriminating fig varieties and
accessions in collections and their capability to determine the degree of diversity
discovered in a defined geographical region or country. In some studies, several
morphological and chemical traits were identified as the most informative for dis-
criminating fig varieties (Çalişkan & Polat, 2012b; Hssaini et al., 2020a, b).
Çalişkan and Polat (2012a), among 64 IPGRI and CIHEAM fig descriptors,
including plant and fruit characters, identified 37 characters as more beneficial for
separating 76 Turkish fig accessions. Traits that showed higher discrimination
capacity among fig accessions were: (a) for the description of plant and leaf: apical
dominancy, lateral shoot formation, leaf shape, number of lobes, length of the cen-
tral lobe, leaf area, and leaf width; and (b) for a description of fruit: fruit length and
width, pH, fruit flesh color, abscission of the stalk from the twig, fruit neck length,
fruit weight, and antioxidant capacity. The authors highlighted that the phytochemi-
cal properties could also be success criteria for describing fig genetic resources.
Antioxidant capacity, anthocyanins, phenols, glucose, and fructose have been
reported helpful descriptors. These characters are, in fact, economically essential
and contribute to the nutritional value of the fig accessions and thus represent criti-
cal selective markers for the selection of genotypes during the breeding program. In
another study, Çalişkan and Polat (2012b) confirmed the significant influence of
genotype on phytochemical and fruit quality properties. The Turkish variety ‘Bursa
Siyahı’ with dark black fruit skin, for example, has the highest levels of total antho-
cyanins, total phenolics, and antioxidant capacity. It has also been observed that
total phenolics correlate moderately with antioxidant capacity and less with total
anthocyanins. These observations confirm the importance of a comprehensive study
from genotype to phenotype, including pomological and biochemical characters of
individual accession. Biochemical attributes provide additional information about
the nutritional value of a defined accession, and only complete information about
the individual genotype allows the selection of the most appropriate accessions for
cultivation.
Hssaini et al. (2020b), in the analysis of 96 Moroccan and 44 introduced fig vari-
eties, demonstrated that fruit dimensions and skin color coordinates were the most
informative for discrimination of fig accessions. In another study of five Moroccan
and six introduced fig accessions, Hssaini et al. (2020a) also confirmed the effec-
tiveness of combined morphological and biochemical characters in pre-breeding
programs. The most relevant pomological characters were reported fruit as weight,
size, stalk, and neck dimensions, whereas among biochemical descriptors, organic
acids, fructose, lightness (L), Chroma (c*), and free radical scavenging activity
based on ABTS assay were the most informative.
A Brazilian study of 42 fig accessions from the Active Germplasm Bank (AGB)
of Fig belonging to the Faculty of Engineering of Ilha Solteira (FEIS) of the
University of Sao Paulo State, among fig leaf measures studied, identified the
3 Fig Tree Genome and Diversity 47

number of leaf lobes as the most efficient marker for accession discrimination with
high heritability (0.85) (Rodrigues et al., 2019) which is in agreement with Çalişkan
and Polat (2012a). The Iranian study reported that most discriminator parameters
depended on a leaf, fruit size, and growth form (Fatahi et al., 2017). In an extensive
study of Spanish figs, Giraldo et al. (2010) analyzed 134 quality parameters of the
tree, leaf, fruit, and two biological variables of 35 accessions grown in the fig col-
lection in the Finca La Orden-Valdesequera in Badajoz, Spain. Most of the descrip-
tors have been selected from the list of descriptors developed for the common fig by
IPGRI and CIHEAM (2003), and, in addition, 43 new descriptors have been
employed as informative for germplasm evaluation. The aim was to select a subset
of highly informative descriptors that were able to distinguish accessions according
to the importance of: (1) production type; (2) morphology of the main and breba
crop; (3) morphology of five-lobed and three-lobed leaves; (4) morphology of entire
leaves, and (5) tree morphology. They performed PCA analysis and a total of 34
variables were selected and broken down into 97 characters based on the highest
discrimination ability in the first four PCs. The first 11 PCs explained 93.3% of the
total variability, while the first four PCs explained 79%. The authors have shown
that for the generation of representative data with morphological evaluation, the
optimization of the approach is needed, including careful selection and reduction of
the number of morphological parameters. Such optimization prevents redundant
information and contributes to saving time and resources.
Morphological evaluation has also provided information on the diversity of the
common fig. Hssaini et al. (2020a, b) reported that the Moroccan local fig clones are
a rich source of variability. Similar conclusions have been reported for wild Iranian
figs (Fatahi et al., 2017). In contrast, in Brazil, fig accessions showed medium to
high diversity expressed with morphological descriptors and contributed to identi-
fying accessions with attractive agronomic traits with the potential for their use in
further genetic improvement programs (Rodrigues et al., 2019).

3.2 Molecular Evaluation

The development of DNA markers has further enriched the characterization

approaches of fig genetic resources by an additional description of each sample with
the tree’s accurate fingerprint (DNA profile). These markers have complemented
classical, morphological markers traditionally used to evaluate individuals and bio-
chemical markers (allozymes, isozymes), considered the first molecular markers
able to detect amino acid sequence variances (Adhikari et al., 2017). The main fea-
tures of DNA markers are that they are not influenced by environmental conditions
or the developmental stage of individuals and are detectable in all tissues of an
organism. In addition, the variation of alleles makes them a powerful genotyping
tool for the distinction of varieties and assessment of genetic diversity within and
between populations. The potential that DNA markers can be linked to a particular
trait or locus is also used in plant breeding programs for marker-assisted selection.
48 D. Bandelj et al.

They, therefore, support many research areas, from taxonomy, phylogeny, popula-
tion, and ecology genetics to biotechnology, plant breeding, and forensics. However,
DNA markers have their strengths and weaknesses, and their selection depends on
the research’s purpose and informativeness (Riaz et al., 2021).
Most of the studies of the common fig involving DNA markers have aimed to
evaluate the genetic diversity of local germplasm, to establish the identification key
for varieties which allow fast distinguishing of varieties present in a collection/
country/region with a minimum number of DNA markers, and to resolve variety
denomination. The diversity of the common fig has been investigated with several
molecular markers such as RAPD (Random Amplified Polymorphic DNA), AFLP
(Amplified Fragment Length Polymorphism), RAMPO (Randomly Amplified
Microsatellites Polymorphisms), ISSR (Inter-Simple Sequence Repeats), and SSR
(Simple Sequence Repeats or microsatellites). In addition, some genetic studies
have employed different DNA marker systems and presented their comparison and
capability to discriminate among fig varieties (Cabrita et al., 2001; Ali-Shtayeh
et al., 2014).
Dominant, multilocus PCR-based DNA markers such as AFLP (Vos et al., 1995)
and RAPD (Williams et al., 1990; Welsh & McClelland, 1990) amplify more loci in
a single analysis simultaneously, and it is expected that they cover a higher propor-
tion of the genome. The AFLP method detects polymorphisms generated with selec-
tive PCR amplification of previously digested genomic DNA with two restriction
enzymes. This method generates 50–100 fragments per assay, a unique genome
fingerprint under investigation. In contrast, RAPD detects polymorphisms of DNA
fragments generated with short, arbitrary primers in the PCR, resulting from changes
in nucleotide sequences at or between the primer binding sites. The RAPD method
amplifies several loci in a single assay, ranging from 0.5 to 5 kb (Adhikari et al.,
2017). The main feature of these two markers is that they can be used in genetic
studies without any prior knowledge of the DNA sequence and can detect polymor-
phisms between closely related genotypes generating unique DNA fingerprints.
However, their limitation is based on the fact that we cannot distinguish heterozy-
gous from homozygous individuals, and the methods demand strict control of pro-
tocol conditions to ensure the repeatability of the results.
For example, in the common fig, RAPDs combined with pomological markers
have been employed to assess Palestinian ecotypes collected in the ex-situ field gene
bank BERC-Til Botanic Gardens, Biodiversity, and Environmental Research Center
(BERC), Til, Nablus (Ali-Shtayeh et al., 2014). Analysis has demonstrated consid-
erable genetic diversity with the potential to improve the crop. Researchers have
also reported a higher discrimination power of combined pomological and molecu-
lar markers, in contrast to using each marker alone. RAPDs have also been employed
to study the genetic structure of 14 individuals representing cultivated Iranian fig
populations (Zolfaghari et al., 2019). Dominant AFLPs, RAPDs, and biochemical
markers were also used in Turkey to evaluate a clone of a variety ‘Sarizeybek’ and
eleven clones of ‘Sarilop’, recognized as the main commercial variety used for dried
figs in Turkey (Cabrita et al., 2001). Five isoenzyme systems discriminated against
two varieties, while cluster analysis based on RAPDs divided clones into two
3 Fig Tree Genome and Diversity 49

groups, although discrimination among all the clones was not possible. Higher dis-
crimination power of clones was demonstrated with AFLPs, where 8 combinations
of EcoRI/MseI primers clearly distinguished all eleven ‘Sarilop’ clones. The authors
concluded that both RAPD and isozymes are useful for distinguishing varieties.
Although RAPDs demonstrated lower polymorphism between clones, they were
able to discriminate, whereas AFLPs were appropriate markers for discrimination
of closely related clones within the fig variety.
The RAMPO method combines RAPD primer and hybridization of amplified
RAPD fragments with microsatellite probes. Fingerprints that are microsatellite-
based may display polymorphisms (Sharma & Singh, 2021). However, the com-
plexity of banding patterns is low, and the method is susceptible (Adhikari et al.,
2017). Chatti et al. (2008) designed the RAMPO method based on RAPD and ISSR
assays to test the method's capability for differentiation of Tunisian fig varieties.
The method showed a good capability for identity determination and resolved the
mislabel of varieties.
The ISSR method includes amplifying the DNA region positioned between the
two microsatellites with microsatellite-complementary PCR primers that can be
used in an anchored or unanchored version (Sharma & Singh, 2021). The ampli-
cons’ length is usually between 200 and 2000 bp, and their polymorphisms depend
on the abundance and hypervariability of microsatellites in the genome (Adhikari
et al., 2017). The method is usually used for cultivar distinction (Sharma & Singh,
2021), although the dominant inheritance and homoplasy are the main limitations
(Adhikari et al., 2017). ISSR markers have been used to investigate genetic relation-
ships among cultivated and wild figs from Jordan, showed intermediate to high
discriminating power, and thus provided the basic information necessary for con-
serving accessions in gene banks (Almajali et al., 2012).
DNA barcoding is another approach preferentially used to identify plants at the
species level. The method involves the production of PCR amplicons from the par-
ticular regions and their sequencing. For example, in plants DNA regions from the
chloroplast genome (for example, matK, rbcL) and internal transcribed spacer (ITS)
of nuclear ribosomal DNA are usually analyzed (Cheng et al., 2016).
DNA barcoding, involving amplification of the ITS regions, matK and trnH-
psbA, has been used in an attempt to differentiate seven fig accessions (varieties)
from the Centro de Investigaciónen Biotecnología of Instituto Tecnológico de Costa
Rica (Castro et al., 2015). In general, the authors reported low genetic variation
observed between the genotypes, but the matK barcode was a reliable and cost-
efficient alternative for identifying samples. The authors also presented the strengths
and weaknesses of barcodes for F. carica genotype identification at the intraspecific
level, which should be considered before their use.
The potential use of ITS sequences for diversity study was tested in a set of
twelve economically important fig varieties from Tunisia (Baraket et al., 2009). The
variation of ITS sequences allowed the establishment of relationships between
closely related genotypes, indicating ITS as a valuable tool for clarifying relation-
ships at a lower taxonomic level.
50 D. Bandelj et al.

Oliveira et al. (2012), in an attempt to link the genetic profile with the metabo-
lome of the fig variety, analyzed five Portuguese F. carica cultivars with two plastid
regions (trnH-psbA, rbcL), six leaf morphological descriptors, and chemical com-
pounds (phenols, sterols, and triterpenes). As a result, the correlation between
genetics and metabolome has been confirmed for the variety ‘Preta Tradicional’
only. In addition, the chloroplastic rbcL region was reported as potentially useful for
fig authenticity purposes.

3.2.1 Development of Fig Microsatellites

Microsatellites, or simple sequence repeats (SSRs), have an important place in plant

genome research (Baruca Arbeiter et al., 2021) and are believed to be the most pow-
erful technique for identification and variety protection (Sharma & Singh, 2021).
Microsatellite sequence consists of repetitive DNA, made of short motifs usually
long from one to six bp, which is repeated repeatedly. The variable number of
repeats enables accurate genetic profiling of each individual. Microsatellites com-
bine several properties of an ideal marker; they are codominant, highly polymor-
phic, abundant in genome (genomic SSRs) and transcriptome (genic SSRs), their
amplification is fast and easy, alleles are easily scored and interpreted, and genotyp-
ing results are highly reproducible, allowing an easy exchange of data among labo-
ratories (Kumar et al., 2009). Although most of the studies have been focused on the
development of genomic SSRs, genic SSRs usually derived from Expressed
Sequence Tags (ESTs) (EST-SSRs) are also attractive for gene mapping and func-
tional studies (Rudd, 2003; Ozgenturk et al., 2010; Baruca Arbeiter et al., 2017).
They are positioned in expressed sequences, generated with next-generation
sequencing technology, and represent a snapshot of gene expression during a par-
ticular developmental stage and under defined conditions when sampling in cells,
tissues, or organs of an organism (Rudd, 2003; Kalia et al., 2011). Therefore, their
value relies on the potential that they are linked to a defined function or locus of
agronomic interest. The largest set of EST-SSRs has been obtained from the fruit
syconium, leaf, and stem transcriptome (Mori et al., 2017). The authors designed
3467 locus specific primer pairs for EST-SSR markers of which 58 were integrated
into a high-density consensus linkage map.
Genomic microsatellites are routinely and most frequently used for genetic stud-
ies and management of plant genetic resources. However, the successes of genetic
analysis rely on selecting the most informative microsatellite markers that allow the
detection of polymorphisms. The selection of markers should be based on the essen-
tial parameters of diversity, such as the number of amplified alleles (n), effective
alleles (ne), observed (Ho) and expected heterozygosity (He), polymorphic informa-
tion content (PIC), and probability of identity (PI). Therefore, the screening process
and quality testing of candidate loci are essential steps in genetic analysis (Selkoe
& Toonen, 2006).
Several microsatellite markers have been published for closely related species
such as Morus sp. and Ficus sp., but their informativeness for diversity studies and
transferability to the common fig should be first tested. The homologous DNA
3 Fig Tree Genome and Diversity 51

sequences in microsatellite flanking regions and evolution distance of species may

influence the transferability (Knap et al., 2016). Only three sets of genomic micro-
satellite markers have been developed and fully characterized for common fig diver-
sity studies.
The first set of microsatellites, named MFC, was released by French researchers
in 2001 (Khadari et al., 2001). From 20 developed locus-specific primer pairs, 8
produced polymorphic alleles in 14 varieties and two wild populations, sampled in
the fig germplasm collection of the Conservatoire Botanique National Méditerranéen
de Porquerolles, France. The MFC set amplified from 2 to 6 alleles, with 4.25 alleles
per locus. The highest number of alleles was obtained with MFC1 (6 alleles) and
MFC2 (5 alleles), and others amplified 4 or 3 alleles. The authors have demon-
strated the transferability of MFC microsatellites to other 17 Ficus species.
Giraldo et al. (2005), a research team from Spain, developed the second, LMFC,
set of fig microsatellites. They obtained 26 polymorphic microsatellites from a
genomic library enriched for CT/AG repeats. The set of microsatellite markers was
characterized in a group of 15 fig varieties from Spain and France. The highest
power for distinguishing fig varieties has been demonstrated with locus LMFC30,
with which 8 alleles have been amplified, and the lowest PI was obtained (0.15).
Comparison of DNA profiles allowed discrimination of nine varieties out of 15. Six
samples (two groups of three varieties) remained undistinguished and showed iden-
tical allelic profiles. Molecular data corresponded with morphological evaluation
and confirmed the inconsistency in the naming of fig varieties. The authors assigned
the low values of variability parameters and the lack of clear geographical group-
ings to a narrow genetic base in the cultivated common fig. Giraldo et al. (2005)
confirmed the transferability of some LMFC microsatellite markers to another 13
Ficus species and Morus alba L. and Morus nigra L. species.
The limited number of highly polymorphic microsatellite markers in the com-
mon fig encouraged a Slovenian research group to develop another set of microsat-
ellites. Fifteen FCUP microsatellites were released in 2007 (Bandelj et al., 2007)
and characterized 22 wild and 19 cultivated fig samples. The highest number of
amplified alleles in cultivated fig varieties was obtained at loci FCUP8 (7 alleles),
FCUP68 (6 alleles), FCUP70 (5 alleles), and FCUP44 (5 alleles). Nine out of 15
loci had PIC higher than 0.5, classifying loci as informative markers. The lowest PI
was detected at locus FCUP8, FCUP44, and FCUP68, indicating that loci are appro-
priate for genotype identification. An additional sixteen primer pairs of the FCUP
series were published in 2016 (Knap et al., 2016). These markers genotyped 18 fig
varieties and 65 wild samples from Slovenia and Croatia. Twelve loci were informa-
tive with PIC higher than 0.5. The highest number of polymorphic alleles was
detected with FCUP40 and FCUP95 (n = 8), 7 alleles were amplified with FCUP1
and FCUP3, locus FCUP24 amplified 6 alleles, and five polymorphic alleles were
found on loci FCUP17, FCUP18, FCUP26, FCUP28, FCUP59, FCUP75, FCUP81,
and FCUP91. These markers demonstrated high discrimination power and represent
an efficient tool for verifying the origin of seedling material in nurseries, distin-
guishing closely related varieties, and managing fig genetic resources collections.
All markers were also successfully cross-amplified in five closely related species
52 D. Bandelj et al.

belonging to the Moraceae family, namely Ficus (F. barteri Sprague, F. rubiginosa
Desf. ex Vent., F. benjamina L.) and Morus (M. alba L. and M. nigra L.).

3.3 Application of Microsatellites in Fig Diversity Studies

Microsatellites of the common fig have been used in many fig producing countries,
mainly for evaluation of genetic diversity of fig varieties and other accessions pres-
ent in ex-situ collections or maintained in-situ at growing sites, for the establishment
of the identification key of varieties, and to assess the diversity of caprifigs
(Table 3.1). Knowledge of the genetic diversity range represents a valuable contri-
bution to the optimization of fig tree breeding programs (Ganopoulos et al., 2015;
Essid et al., 2015) and the conservation and improvement of local fig germplasm
(Ben Abdelkrim et al., 2015; Baraket et al., 2011; Saddoud et al., 2011; Essid et al.,
2015). Interlaboratory comparison of fig varieties' genotyping profiles with micro-
satellites has also been shown (Hladnik et al., 2018). Some studies compared differ-
ent marker systems and presented their potential for discrimination of fig accessions
(Ikegami et al., 2009; Chatti et al., 2010; Ikten et al., 2010; Baraket et al., 2011;
Abou-Ellail et al., 2014).
Microsatellites have been used to characterize the Moroccan fig germplasm col-
lection managed by the Institut National de la Recherche Agronomique (INRA)
Meknes. Achtak et al. (2009) genotyped 75 accessions, including eight caprifigs, 51
local accessions, 11 foreign accessions, and five accessions of unknown origin.
Among 17 microsatellite loci used in the study, two optimal combinations of primer
pairs have been proposed to discriminate all accessions. The combination LMFC3
0 + MFC2 + MFC3 + FSYC01 + MFC9 allowed almost all analyzed samples to be
identified except for two genotypes, which were further differentiated by adding the
locus MFC11 to the combination. The lowest PI values have been reported for loci
LMFC30, MFC2, MFC3, and MFC1 (PI < 0.2), which confirms the good ability of
loci to distinguish varieties included in the analysis. In addition, with the help of
microsatellites, cases of synonyms and homonyms, and mislabels in the collection
have been resolved. The study of the Moroccan fig collection was further upgraded
with microsatellite profiling of 277 individual fig trees from 40 sites distributed in 6
geographical regions (Achtak et al., 2010), representing traditional Moroccan agro-
ecosystems. With the same set of 17 microsatellite primer pairs, the highest dis-
crimination power has been noted for loci LMFC30, MFC1, MFC2, and MFC3
(D > 0.80). The extensive study has demonstrated that traditional Moroccan agro-
ecosystems represent active varietal and genetic diversity incubators and are an
important repository of local varieties. Although the varieties are clonally main-
tained, small differences from 1 to 3 alleles among the trees were assigned to
somatic mutations accumulated through the long period of variety cultivation. The
higher number of allele differences among genotypes (4–34 alleles) has been
explained by the sexual reproduction of cultivated figs and wild growing figs in the
3 Fig Tree Genome and Diversity 53

Table 3.1 The list of published scientific articles involving the common fig SSRs in diversity studies
Common fig Fig genetic material
Country SSRs (set) analysed (accessions) Research aims Reference
Algeria 24 SSRs 34 varieties Fig diversity and Boudchicha
(LMFC, denomination et al. (2018)
FCUP)
Brazil 5 SSRs 11 varieties (collection) Fig evaluation and do Val et al.
(MFC) identification (2013)
France SSRs (MFC) 14 varieties and 2 wild SSRs development and Khadari et al.
populations characterization: 20 (2001)
primers
Greece 7 SSRs 90 varieties from different Fig diversity, genetic Ganopoulos
(LMFC, countries (collection) differentiation et al. (2015)
MFC)
Italy 7 SSRs 79 parthenocarpic Fig diversity and Rodolfi et al.
(LMFC, accessions (collection) denomination, creation (2018)
MFC) of fig database
Italy 18 SSRs 181 traditional varieties Fig diversity and Costa et al.
(LMFC, denomination (2017)
MFC, FCUP)
Morocco 17 SSRs 277 fig trees from 6 Fig diversity in Achtak et al.
(LMFC, regions traditional (2010)
MFC) agroecosystems
Morocco 17 SSRs 8 caprifigs, 51 local,11 Fig diversity and Achtak et al.
(LMFC, introduced, 5 unknown denomination (2009)
MFC) accessions (collection)
Slovenia 12 SSRs 23 Adriatic varieties, 218 Fig diversity and Knap et al.
(FCUP, accessions from NCGR genetic relationships (2018)
LMFC) collection
Slovenia SSRs 18 varieties and 65 wild SSRs development and Knap et al.
(FCUP) forms characterization: 16 (2016)
primers
Slovenia SSRs 19 varieties and 22 wild SSRs development and Bandelj et al.
(FCUP) forms characterization: 15 (2007)
primers
Slovenia 6 SSRs 90 fig accessions (84 Comparison of the Hladnik et al.
& France (LMFC, from the East Adriatic, 6 genotyping results of (2018)
MFC, from the French two laboratories
FSYC01) collection)
Spain 9 SSRs 229 accessions from Establishment of core Balas et al.
(LMFC, Spain and abroad collection (2014)
MFC)
Spain 21 SSRs 42 local accessions Genetic relationships Perez-Jiménez
(LMFC, belonging to 12 local among fig accessions et al. (2012)
MFC) denominations and two
caprifigs
(continued)
54 D. Bandelj et al.

Table 3.1 (continued)

Common fig Fig genetic material
Country SSRs (set) analysed (accessions) Research aims Reference
Spain 11 SSRs 196 samples from Establishment of a Giraldo et al.
(LMFC, different growing areas in genotyping database of (2008)
MFC) Spain and 13 samples reference profiles
from abroad
Spain SSRs 15 varieties from Spain SSRs development and Giraldo et al.
(LMFC) and France characterization: 26 (2005)
primers
Tunisia 13 SSRs 30 cultivated accessions Differentiation of Essid et al.
(LMFC, accessions (2021)
MFC)
Tunisia 17 SSRs 19 varieties, 52 wild figs Identification, diversity Ben
(LMFC, from 4 locations study Abdelkrim
MFC) et al. (2015)
Tunisia 13 SSRs 20 caprifigs Differentiation of Essid et al.
(LMFC, caprifigs, fig diversity (2015)
MFC)
Tunisia 6 SSRs 72 local ecotypes Identification Saddoud et al.
(MFC) (2007)
Turkey 15 SSRs 96 native caprifigs Genetic population Çalişkan et al.
(LMFC, structure (2018)
MFC, FCUP)
USA 15 SSRs 194 accessions from Fig diversity and Aradhya et al.
(LMFC, different countries genetic relationships (2010)
MFC) (NCGR collection)

agroecosystem. This spontaneous hybridization represents an important driving

force for developing varieties with new qualitative properties.
Severe genetic erosion of local fig germplasm and the confusion in the denomi-
nation of varieties in Tunisia have encouraged the evaluation programs of fig genetic
patrimony (Saddoud et al., 2007). The first attempt at the preservation of fig eco-
types included the inventory and propagation of cuttings of around 100 ecotypes
that are ex-situ maintained in different collections (Institut Supérieur Agronomique
(ISA), Chott Meriem; Institut des Régions Arides (IRA), Medenine; Centre de
Recherches Phoénicicoles (CRPh), Degache; Commissariat Régional au
Développement Agricole (CRDA), Gafsa; Groupement Interprofessionnel des
Fruits (GIF), Sbikha) or are in-situ preserved (Saddoud et al., 2007). Six MFC
markers have been used for genotyping 72 local ecotypes originating from different
Tunisian areas. Fingerprinting allowed the establishment of the first identification
key for Tunisian varieties.
Ben Abdelkrim et al. (2015) performed genotyping of Tunisian fig germplasm,
including 19 fig varieties and 52 wild fig trees from four different locations with the
identical primer pairs set of 17 microsatellites as previously reported in an analysis
of Moroccan fig germplasm (Achtak et al., 2009, 2010). On average, the number of
amplified alleles per locus was similar to that reported in other fig diversity studies
3 Fig Tree Genome and Diversity 55

(4.35 alleles/locus). In Tunisian fig samples, the highest PIC was found in loci
MFC3, LMFC30, and MFC11. These three loci, together with MFC19, constitute
the identification key for Tunisian fig germplasm. Although some of the samples
were not discriminated with this combination, the authors found other loci that
could distinguish these genotypes. This result suggests the careful selection of mic-
rosatellite loci for identification purposes and the narrow genetic base of some fig
genotypes. The newest study, conducted by Essid et al. (2021), employed 13 micro-
satellites LMFC and MFC loci and presented their characteristics in 30 cultivated
Tunisian fig accessions. PI values of loci were relatively high, indicating a lower
capability to differentiate accessions under evaluation. Similar results were reported
in Tunisian caprifigs from different geographic regions showing distinct phenotypic
traits (Essid et al., 2015). The same set of 13 microsatellite LMFC and MFC loci
generated a low number of alleles (2 or 3 alleles per locus), and diversity parameters
were below the range observed in other studies. Although the authors attributed this
result to a low genetic diversity of the Tunisian male trees, the limited number of
analyzed caprifig trees (20) and the selection of loci with lower diversity informa-
tion content could have influenced the final observations. The authors highlighted
the need for additional studies of caprifigs to obtain a better insight into their genetic
diversity.
The study of the genetic population structure of caprifigs from Turkey showed
the opposite result analyzing 96 native caprifigs with 15 loci of series MFC, LMFC,
and FCUP, which were chosen based on their high PIC values (Çalişkan et al.,
2018). On average, 8.3 alleles were amplified, ranging from 3 (LMFC18, LMFC23)
to 14 alleles (FCUP38, FCUP8). The study showed a great genetic diversity of cap-
rifigs assigned to generative propagation with seeds in nature. The population
genetic structure analysis of caprifigs demonstrated grouping according to the geo-
graphical region of sampling.
The presence of figs from ancient times in Italian territory has been supported
with archaeobotanical and historical evidence, and the long tradition of cultivation
has contributed to a large number of varieties in the country. Rodolfi et al. (2018)
genotyped 79 parthenocarpic fig accessions from a private, ex-situ collection in
Pescia (Tuscany, IT) with MFC and LMFC microsatellites. The analysis demon-
strated the existence of 56 different genotypes and has contributed to resolving
cases of homonyms, synonyms, and mistaken denominations. As a result, a data-
bank of Tuscan and other Italian fig varieties has been created. In another study of
Italian fig germplasm, Costa et al. (2017) employed 18 FCUP, LMFC, and MFC
microsatellites and presented their characterization in 181 traditional fig varieties
and genotypes cultivated in Apulia, Basilicata, Calabria, Campania, and Sicily
(south Italy). Nine out of 18 loci displayed the PIC value higher than 0.5; the most
informative markers were FCUP27, FCUP38, MFC1, MFC2, MFC3, MFC11
(PIC > 0.6), LMFC28, MFC7 (PIC > 0.5) and LMFC30 (PIC > 0.7). Their study
demonstrated the richness of the fig genetic resources in Southern Italy and helped
resolve fig varieties denomination.
In Spain, Giraldo et al. (2008) analyzed 196 samples from different growing
areas and 13 samples from the USA, France, Israel, Portugal, and Turkey. Among
56 D. Bandelj et al.

20 polymorphic SSR markers (series LMFC, MFC) tested, eleven amplified one
locus, and these markers were further characterized for their polymorphisms.
Microsatellite data helped establish the database with DNA profiles of 104 refer-
ence genotypes from 209 starting accessions. The redundancy of samples in a germ-
plasm collection is not desirable, and before incorporating new accessions into a
collection, it is worth testing their identity. The study demonstrated that microsatel-
lites are an excellent tool for collection management and can identify synonyms and
homonyms. The applicability of microsatellites for the evaluation of local fig germ-
plasm has been demonstrated by Perez-Jiménez et al. (2012). In their study, 42 trees
belonging to 12 local denominations and two caprifigs were genotyped with 21
LMFC and MFC microsatellite primers. Their allelic profiles were compared with
15 reference varieties from collections of the Conservatoire Botanique National
Méditerranéen de Porquerolles (France) and Estación Experimental Agraria de
Elche (Spain), and a local nursery from Córdoba (Spain). The highest PIC (higher
than 0.5) was obtained with loci MFC2, MFC3, MFC6, LMFC12, LMFC13,
LMFC30, and LMFC38. The clustering method separated the studied Andalusian
fig trees into 11 distinct groups corresponding to their local denominations. These
data will serve for fig germplasm conservation strategy in Spain.
The National Clonal Germplasm Repository (NCGR) at Davis, California (USA)
is one of the largest collections of fruit and nut crops that collect, preserve, evaluate,
and distribute the genetic resources of crops for current and future needs. The col-
lection is home to numerous fig accessions from all over the world. Aradhya et al.
(2010) have analyzed 194 germplasm accessions representing all four types of figs
with fifteen MFC and LMFC loci. Cluster analysis based on microsatellite profiles
revealed ten groups, which have been confirmed with PCA. With PCA, more dif-
ferentiation has been noted in some groups. The gene diversity was higher within
the groups than among the groups. The study's main conclusion is that fig germ-
plasm possesses substantial genetic polymorphism but exhibits narrow differentia-
tion. In addition, Turkmenistan figs showed different genetic backgrounds compared
to Mediterranean and Caucasian fig germplasm.
Knap et al. (2018) have compared genotyping profiles of 23 Adriatic fig varieties
with 218 accessions from the Californian NCGR collection obtained with 12 FCUP
and LMFC SSR loci. In this study, the number of amplified alleles per locus ranged
from 4 to 14. The average number of amplified alleles was relatively high (8.33)
compared with other fig diversity studies. In addition, seven loci out of twelve had
a PIC value higher than 0.5, indicating the high polymorphic information content of
the loci used. With eight loci, it was possible to discriminate 166 genotypes. The
analysis confirmed significant genetic diversity in the common fig, attributed to the
careful selection of highly informative SSR markers and large sample size with
redundant worldwide accessions. The important achievement of the study was the
identification of several international synonyms of varieties expanded worldwide,
originating from the United States of America (‘Ventura’, ‘Calvert’, ‘Figue D’Or’,
‘Capitola Long’), Georgia (‘Asashendu green figs’, ‘Shilda Purple-Bronze Fig’,
‘GIHVO Italian Green Fig’), Azerbaijan (‘Belaya Fraga’, ‘Blanche Fig’) and Turkey
(‘Patlican fig’). In addition, 17 out of 23 North Adriatic varieties had distinct
3 Fig Tree Genome and Diversity 57

genotypes compared to the NCGR collection, indicating that fig varietal structure
and diversity in Slovenia is unique and represents an important reservoir of genetic
variation for cultivated figs. Local varieties with unique genotypes can significantly
promote the region and produce valuable local products.
Algerian cultivation of figs provides basic income in the rural and marginal
areas, and the country is ranked third among the most important producing coun-
tries of figs in the world, after Turkey and Egypt (Boudchicha et al., 2018). Despite
the importance of its cultivation, genetic erosion has been reported, and the Algerian
Ministry of Agriculture launched a developmental program to improve fig produc-
tion in the country. Boudchicha et al. (2018) presented the first molecular report of
fig diversity. The authors evaluated 34 fig varieties of San Pedro, Smyrna, and com-
mon types from ex-situ and in situ with 24 microsatellite primers. With the highest
PIC, five markers (LMFC30, FCUP038, FS4-11, FCUP044, FCUP070, and
FM4-11) showed an excellent discriminatory capacity for Algerian fig varieties.
Genotyping revealed cases of synonyms, homonyms, and mislabelling errors in the
collections and demonstrated the significant genetic variability with a narrow
genetic structure.
In Brazil, the cultivation of the common fig is increasing due to the growing
demand for its fruits in the food industry (do Val et al., 2013). To overcome some
pest and disease problems related to the cultivation of a single variety, ‘Roxo de
Valinhos’, and to diversify cultivation with other fig varieties, an evaluation of
eleven varieties maintained in the germplasm bank located in Lavras, Minas Gerais
has been investigated by the use of five MFC loci. The allelic variation allowed the
identification of 10 genotypes and two synonymous individuals. In this study, the
lowest PI values were obtained for markers MFC1, MFC3, and MFC7, indicating
good discriminatory capability for the Brazilian collection.
In Greece, seven LMFC and MFC microsatellites have been used to characterize
ninety varieties from Greece, Italy, Cyprus, Spain, Turkey, and France in an ex-situ
collection (Ganopoulos et al., 2015). The diversity parameters of all selected mark-
ers were highly informative, with PIC higher than 0.5. Although substantial genetic
polymorphism has been reported, limited genetic differentiation among the differ-
ent geographical groups has been noted, probably due to the common origin of
analyzed varieties. For example, fig genotypes from Cyprus were genetically differ-
ent from the rest of the accessions.
Microsatellites are confirmed as a very useful tool for collection management.
The high costs of managing a large number of accessions in the collection, primarily
due to redundancies and duplications, led to the search for new solutions to reduce
accessions without the decline in the overall diversity of the plant. One of the
approaches includes creating a core collection composed of a reference subset of
genotypes selected from a larger set of varieties. Balas et al. (2014) found that the
maximum diversity could be achieved with a maximization strategy (M strategy)
based on microsatellite data. Their study was conducted using nine LMFC and
MFC microsatellites on 229 fig accessions from Spain and abroad, maintained at
Centro de Investigación La Orden fig germplasm bank (Spain). M strategy maxi-
mizes the number of alleles sampled at each locus, and an iterative process
58 D. Bandelj et al.

eliminates the redundant accessions. It was found that with the subset of 30 fig
accessions (only 13% of the entire collection), it is possible to cover all the SSR
alleles. In addition, in comparing allele frequency distribution, He and HO showed
no significant differences between the whole and core collection. M strategy seems
to be an efficient method for reducing fig accessions in the collection. The main
consideration is that some locally spread fig ecotypes with a rare combination of
alleles could be lost with this strategy; thus, they require special attention for con-
servation. According to Balas et al. (2014), economically important varieties can be
additionally included in the core collection.
Some genetic studies of figs have employed different marker systems for diver-
sity or identification proposes. Chatti et al. (2010) compared the ability of RAPD,
ISSR, RAMPO, and SSR to detect polymorphism and to establish the genetic rela-
tionships among fig varieties. A comparison of the number of polymorphic markers
obtained with RAPD, ISSR, and RAMPO demonstrated a similar and efficient
capability of polymorphism detection. SSR analysis employed six MFC loci, of
which five were able to discriminate fig varieties except for Kahli1 and Soltani2
samples. These two samples were distinguished with combined RAPD, ISSR, and
RAMPO markers. This result is expected and can be assigned to the nature of the
aforementioned markers, which cover a few random regions in a single PCR reac-
tion, whereas one microsatellite locus covers a very small part of the genome.
Difficulties with discriminating samples with SSRs were already reported by Knap
et al. (2017), when the set of seven FCUP loci was not able to discriminate Slovenian
varieties ‘Zeleni (green) Matalon’ and ‘Črni (black) Matalon’ despite their apparent
differences in the color of the fruit peel. However, these two varieties were previ-
ously successfully distinguished by selecting other combinations of loci (Bandelj
et al., 2008). This result additionally confirmed that the selection of markers for
identification procedures is crucial, and besides the informativeness of markers, it
depends on the genetic background and closes the genetic relationship of the plant
material under investigation.
Tunisian fig germplasm was evaluated through the comparison of AFLPs and
SSRs. Baraket et al. (2011) genotyped 38 accessions, including 30 varieties and
eight male trees from different regions. This study used six MFC microsatellites and
six EcoRI/MseI primer pairs. Both marker systems showed a large genetic diversity
of Tunisian figs and gave good and comparable results, although their nature dif-
fered. According to the authors’ opinion, the advantage of AFLPs is a high effective
multiplex ratio value (the number of bands analyzed simultaneously per experi-
ment) and estimated marker index (the amount of information obtained per experi-
ment for a given marker system). Furthermore, varieties were clustered independently
from their geographical origin, horticultural classifications, and tree sex phenotype
with both markers.
In Egypt, some local fig names have been generated by farmers based on fruit
color and flavor, and the presence of homonyms and synonyms characterizes the
local germplasm. Abou-Ellail et al. (2014) used two biochemical markers, isozymes
3 Fig Tree Genome and Diversity 59

Polyphenol Oxidase, Peroxidase, and 6 SSR (MFC) loci, to establish the molecular
key for fig identification to provide necessary information for breeding programs
and conservation of local germplasm. The analysis was performed on 7 fig varieties
sampled in different growing areas. The study has demonstrated that isozymes are
suitable, but SSRs were more efficient for genetic variability and fig cultivars
fingerprinting.
Ikegami et al. (2009) analyzed the genetic diversity of 8 varieties from Japan, 4
from the USA, 5 from France, and 2 from Spain with ISSR (13 primer combina-
tions), RAPD (19 primers), and SSR (13 loci) markers. Examination of each marker
system demonstrated differences in the separation of samples according to genetic
relationships, and a weak correlation among marker systems was detected. The
study’s main conclusions were that the genetic diversity of analyzed fig samples
was low and that the multiple marker utilization is critical to estimating the genetic
relationships at a varietal level.

4 Gene, Genomic and Transcriptomic Studies of Figs

Technologies for high-throughput sequencing (HTS) have enabled the generation of

large amounts of data compared to technologies based on Sanger’s method.
Sequencing DNA or RNA offers deeper insight into the genome structure and iden-
tification of new genes and mechanisms involved in gene expression. The first high-
throughput sequencing technologies are classified as next-generation sequencing
technologies (NGS), e.g., Illumina and Ion Torrent, whereas the most recent are
designated as third-generation sequencing technologies (TGS) (van Dijk et al., 2018)⁠.
The common fig has received significant attention for analysis with HTS tech-
nologies, which several published scientific articles have confirmed in recent years.
‘Short read’ sequencing was applied for genomic sequences and transcriptome
(reviewed in the next chapters), whereas ‘long read’ sequencing technology was
recently used for genome sequencing (Usai et al., 2020)⁠.
The ‘short read’ NGS technologies involve massively parallel sequencing of
short reads, meaning that millions of individual sequencing reactions occur in paral-
lel (Hu et al., 2021)⁠. However, reconstructing regions with repetitive elements and
identifying large structural variants is often difficult with short reads. The TGS tech-
nologies, on the other hand, including ‘single-molecule real-time’ (SMRT) sequenc-
ing technology developed by Pacific Biosciences (PacBio) and Oxford Nanopore
Technologies (ONT), can sequence longer fragments which leads to superior per-
formance in the analysis of repeated regions, structural variants, haplotype phasing,
transcriptome analysis, and other applications such as closing the gaps of previous
short-read assemblies (van Dijk et al., 2018)⁠.
60 D. Bandelj et al.

4.1 Genomic Studies

4.1.1 Genome Characterization

The common fig is a diploid species with 2n = 26 chromosomes (Flaishman et al.,

2008)⁠. The estimated genome size with flow cytometry is 356 Mbp (Loureiro et al.,
2007)⁠. The first drafts of the common fig genome sequence were generated by Mori
et al. (2017)⁠ and Barghini et al. (2017)⁠ using NGS sequencing technologies. Mori
et al. (2017)⁠ sequenced DNA of the traditional Japanese variety ‘Horaishi’, whereas
Barghini et al. (2017)⁠ sequenced DNA of an old Italian variety ‘Dottato’. Both vari-
eties are classified as common-type figs.
Recently, the genome of the variety ‘Dottato’ was sequenced with PacBio SMRT
technology (Usai et al., 2020)⁠, which enabled the better reconstruction of the
genome, especially parts represented by repetitive elements. The final length of the
assembled genome sequence was ~333 Mbp, corresponding to ~95% of the esti-
mated size. A total of 37,840 protein-coding genes were identified, and repeated
represented 37.39% of the genome assembly. The most abundant repeats were
transposable elements (TE), covering 33.5% of the assembly. Tandem repeats rep-
resent 3.82% of the assembly. Long terminal repeat retrotransposons (LTR-REs)
were the most abundant among the transposable elements, representing 28.03% of
the genome assembly. Altogether, 1867 LTR-REs were revealed (Vangelisti et al.,
2021)⁠. Gypsy elements (62.5%) and Copia elements (33.3%) were annotated, and
the rest of the full-length LTR-REs remained unknown.

4.1.2 New Research Approaches and Findings Based

on the Available Genomes

The available genomes have opened the possibilities for new research approaches,
like genome-wide gene identification and characterization (Song et al., 2021)⁠, iden-
tification of genome-wide single nucleotide polymorphism (SNP) markers, and
generation of high-density genetic maps (Mori et al., 2017)⁠. In addition, available
genomes are an essential platform for transcriptome studies.
Mori et al. (2017)⁠ performed a genome-wide association study (GWAS) and
identified two missense mutations in the RESPONSIVE-TO-ANTAGONIST1 (RAN1)
gene, which were highly associated with sex phenotypes of the investigated figs.
Several analyses were performed to obtain this result. Genome-wide SNPs and
indels polymorphisms were identified based on the ‘Horaishi’ variety draft genome
and genome sequences of five different varieties. Restriction site-associated DNA
sequencing (RAD-seq) analyses were performed on plants of the ‘Horaishi’ and
Caprifig6085 which were cross-pollinated, on its F1 population, and additional
plants with known sex phenotype (22 males, 97 females) and on 3 individuals with
unknown sex phenotype for genotyping and the construction of the high-density
3 Fig Tree Genome and Diversity 61

genetic map. RAN1 is involved in the transduction of ethylene, and a suggested

mechanism is that male plants with normal RAN1 have active ethylene receptors
resulting in the formation of stamens, whereas, in the females, these receptors are
nonfunctional. The role of phytohormones in the differentiation of floral organs was
observed in other plant species (Feng et al., 2020)⁠. Kompetitive allele specific PCR
(KASP) and cleaved amplified polymorphic sequence (CAPS) markers were devel-
oped to detect A or G alleles in the RAN1 gene (G/A in males, G/G in females) or
all three codons that could result in a combination of neighboring SNP, respectively
(Ikegami et al., 2021; Mori et al., 2017)⁠. Surprisingly, segregation distortion was
observed in a specific parental cross on this locus, resulting in a highly male-biased
segregation ratio. Several candidate genes possibly responsible for the segregation
distortion were identified (Ikegami et al., 2021)⁠. Recently, three AGAMOUS genes
were recruited from the F. carica genome. FcAG2-Gall_Stamen and FcAG3-Gall_
Stamen were male-specific (they had 15 bp in the 7th exon absent in FcAG-Pistil)
and were only expressed in the gall and stamen of the male tree fig fruit. A method
for detecting male plants with PCR was developed (AG-Marker) (Wang et al.,
2021b)⁠. The same authors also designed the marker based on 6 SNPs of the RAN1
gene, and both markers were confirmed as an accurate approach for sex
determination.
Publicly available genomes are an essential base for genome-wide gene identifi-
cation and characterization. Song et al. (2021)⁠ studied genes from the basic helix-
loop-helix (bHLH) transcription factor family involved in the anthocyanin
biosynthesis. In total, 118 FcbHLH genes were identified in reference genomes.
Fifteen FcbHLH genes showed increased expression in the peel and female flower
tissues at different main crop developmental stages. The common fig varieties
‘Purple-Peel’ (‘Zibao’) and ‘Green-Peel’ (‘Qingpi’) were used. The ‘Purple-Peel’
variety is a bud mutation of ‘Green-Peel’, a major common fig variety in China
(Wang et al., 2017)⁠. The gene FcbHLH42 synchronicity between expression and the
corresponding anthocyanin content in the female flower and peel was observed. The
impact of the FcbHLH42 gene on anthocyanin accumulation was also demonstrated
with transient expression in tobacco leaves.
Due to the relatively small genome size, the common fig has become a new
model species for fruit trees, and recently methylation-based studies have been per-
formed (Usai et al., 2021)⁠. The study aimed to analyze the pattern of unconventional
epigenetic changes, including N4-methylcytosine (4mC) and N6-methyladenine
(6mA) modifications within the transposable elements (TE) and the impact of TE
methylation on the expression of the neighboring genes. However, an additional
analysis should be implemented to fully elucidate the role of TE methylation pat-
terns on the neighboring gene expression (Usai et al., 2021)⁠.
62 D. Bandelj et al.

4.2 Transcriptomic and Gene-Based Studies

4.2.1 Mechanisms Related to Fruit Development and Ripening

To elucidate biological mechanisms related to plant and fruit development, several

transcriptomes were sequenced (Chai et al., 2017, 2018, 2019; Cui et al., 2019,
2020; Freiman et al., 2014; Ikegami et al., 2013a; Li et al., 2020; Marcotuli et al.,
2020; Qiao et al., 2021; Rosianski et al., 2016a; Wang et al., 2017, 2019)⁠.
The first transcriptome dataset for multiple developmental stages of common fig
fruit was provided by Freiman et al. (2014)⁠. They sequenced RNA obtained from
the red flash variety fruits, classified as common type biferous fig, ‘Brown Turkey’.
As a result, the differential activity of several metabolic processes was observed,
and new ethylene-synthesis genes and the involvement of several transcription fac-
tors during the ripening process were determined.
In order to identify genes involved in fruit-quality formation control transcrip-
tome analysis between receptacle tissues of commercial ripe (first half of stage III)
and tree-ripe (late-stage III) fruits were performed by Cui et al. (2019)⁠. Differentially
regulated genes related to ficins, sugar accumulation, cell-wall modification, ethyl-
ene pathway, and other biological processes involved in fruit quality and ripening
were observed. In addition, genes involved in cell wall modification and their
expression during ripening were studied by Owino et al. (2004), and it was sug-
gested that they are coordinated in time and their intensity of expression to achieve
fruit softening.
To discover the key candidate genes in ripening regulation to achieve more
simultaneous ripening of all fruits on a tree, consequently shortening the harvest
season and making it less labor-intensive, an experiment was performed with
ethephon-treated fruits (Cui et al., 2020)⁠. Ethephon stimulates fig fruits’ growth and
ripening. It was observed that ethephon mainly regulates genes related to the expres-
sion of plant hormones, cell-wall modification, and sugar accumulation in female
flowers and receptacles.
Since fruits harvested before ripening onset will not ripen postharvest, Freiman
et al. (2015)⁠ studied the influence of 1-methylcyclopropene (1-MCP) (an inhibitor
of ethylene), which improves fruit-storage abilities after treatment of fruits on trees,
on ethylene production, and other mechanisms related to fruit ripening. The behav-
iour of genes involved in fruit ripening and genes involved in the prolonged storage
abilities after treatment with 1-MCP was characterized. In addition, genes involved
in the ethylene synthesis were isolated, and their expression pattern during ripening
was also studied by Owino et al. (2006)⁠. In addition, it has been shown that on-tree
abscisic acid (ABA) application synchronizes fruit ripening and maintains its qual-
ity (Lama et al., 2019)⁠, and a model for the role of the ABA-ethylene interaction
during fruit ripening was proposed by Qiao et al. (2021)⁠.
Flaishman et al. (2020)⁠ proposed ABA and ethylene crucial biosynthetic genes,
FcNCED2 and FcACS4, as candidate genes for genome editing to alter the ripening
process of fruits. An attempt to edit the FcNCED2 gene was already performed,
3 Fig Tree Genome and Diversity 63

using Agrobacterium-mediated transformation of a construct carrying the CRISPR/

Cas9 complex components. However, the desired mutation has to be confirmed.
For different common type varieties, it was observed that pollinated fruits had
larger diameters, weight, improved firmness, and more commercially desirable
characteristics compared to the parthenocarpic fruit. This was observed for the
‘Brown Turkey’ variety (Rosianski et al., 2016b)⁠ and recently for fruits of the
‘Dottato’ variety (Marcotuli et al., 2020)⁠. To study gene expression during fruit
ripening in pollinated and parthenocarpic fruits, a transcriptome comparison of
‘Brown Turkey’ fruits was performed by Rosianski et al. (2016a)⁠. Results indicate
that the reproductive part of the syconium coordinates fruit ripening, and the differ-
ences in ABA production between pollinated and parthenocarpic fruit might be the
key to their different ripening characteristics.

4.2.2 Secondary Metabolites

Transcriptome studies related to the flavonoid biosynthetic pathways were con-

ducted using fruit peels (Li et al., 2020; Wang et al., 2017)⁠. Wang et al. (2017)⁠
analyzed ‘Green Peel’ and its color mutant ‘Purple Peel’. Significant and simultane-
ous upregulation was revealed in almost all of the flavonoid and anthocyanin path-
way components and relevant transcription factors in the mature stage of ‘Purple
Peel’ fruits. Li et al. (2020)⁠ used plant material of the variety ‘Bojihong’ and com-
pared transcriptomes at the yellow and red stages. Differentially expressed
anthocyanin-related genes were observed. Three of them (FcCHS1, FcCHI1, and
FcDFR1) and two genes from the R2R3 MYB family of transcription factors
(FcMYB21 and FcMYB123) were additionally characterized after cloning. The reg-
ulatory role in the anthocyanin accumulation of the last two genes was confirmed
with transient expression in apple fruits and calli. In addition, a full-length MADS-
box gene, FcMADS9, significantly up-regulated during the pigmentation process,
was identified (Li et al., 2021)⁠.
A study performed by Wang et al. (2019)⁠ focused on the anthocyanins content
and gene expression in fruit peel and female flower tissue of a variety ‘Zibao’ at
different time points. In addition, treatment with light deprivation was included, and
anthocyanin content in the peel was repressed (suppression of anthocyanin synthe-
sis was also observed on a gene expression level) compared to the control, confirm-
ing the statement that fruit coloration can be firmly and differentially affected by
light. Overall, transcriptome analysis revealed differentially expressed pathways,
specific structural anthocyanin-biosynthesis genes, and several potential transcrip-
tion factors, particularly MYBs. Promoter sequences of FcCHS1 and FcDFR1 were
cloned, and elements related to light response were identified in their promoter
regions. G-box, L-box, Box 4, and GTGA-box were found in the promoter region
of FcCHS1, indicating that FcCHS1 could be regulated by light-induced signal-
transduction elements such as HY5. On the other hand, H-box, I-box, and G-box
elements, required for binding MYB transcription factors, were identified in the
FcDFR1 promoter. The importance of FcHY5 and FcMYB114 in the expression of
64 D. Bandelj et al.

FcCHS1 and FcDFR1 was confirmed with a yeast one-hybrid screen. Recently tran-
scriptome analysis was performed to compare gene expression before and after fruit
bagging of the same variety (Wang et al., 2021a)⁠.
Cao et al. (2016)⁠ studied anthocyanidin synthase (ANS), a key enzyme in antho-
cyanin biosynthesis cloned from syconia (FcANS1). They performed a structural
analysis of the gene and monitored its expression in different fruit tissues. Their
work provided an essential basis for further studies on anthocyanin accumulation
and regulation in fruit. Additionally, it was observed that ABA could initiate antho-
cyanin biosynthesis, which can improve the color and nutritional value of fig fruit
(Lama et al., 2020)⁠.
The first gene of the carotenoid pathway, lycopene beta-cyclase, which is respon-
sible for converting lycopene to beta-carotene, was identified in the common fig,
and a full-length cDNA was cloned by Araya-Garay et al. (2011)⁠. In addition, Zeta
carotene desaturase, involved in carotene desaturation, was also cloned from the
common fig (Araya-Garay et al., 2012)⁠.

4.2.3 Physiological Differences Between Common Fig Types

With the aim to reveal changes in fig fruit physiology of different fig types, Ikegami
et al. (2013a)⁠ sequenced the transcriptome of caprifig (Caprifig 6085) and a com-
mon type variety ‘Houraishi’. Among the differentially expressed genes, chalcone
synthase (CHS) was identified, which was previously involved in the development
of parthenocarpic fruit (Schijlen et al., 2007)⁠.
Fig varieties classified as San Pedro type were also studied to infer the mecha-
nism underlying the development of the parthenocarpic breba and non-parthenocarpic
main crop (Chai et al., 2017)⁠. Fruits of the ‘Asteran’ variety were used for gene
expression studies. Gibberellin (GA)- and auxin-biosynthesis genes were repressed,
and GA and ABA response in main crop female flowers, whereas ABA- and
ethylene-biosynthesis genes were enhanced. The impact of gibberellin and cytoki-
nin treatments on parthenocarpy induction of the main crop was studied on fruits
from the same variety (Chai et al., 2018, 2019)⁠.
Fruit buds developed in the current season can differentiate between developing
the main crop or developing the breba crop in the following year; therefore, the dif-
ferences in gene expression between fruit buds differentiation in common type vari-
ety and San Pedro type variety were studied by Marcotuli et al. (2020)⁠. The two fig
varieties used in the study were the common type variety ‘Dottato’, mainly produc-
ing main crops (rarely brebas), and ‘Petrelli’, belonging to the San Pedro type,
which mainly produces brebas and sometimes a few main crop fruits. Variability in
gene expression patterns was observed between the varieties and between the fruit
buds of the breba and the main crop of the same variety. Particularly in the ‘Petrelli’
main crop, flowering locus T showed a high expression level, whereas in ‘Dottato’,
especially in brebas, the AGAMOUS gene, which is required for normal develop-
ment of stamens and the carpel in flower, was highly expressed compared to the
‘Petrelli’. In addition, several genes related to auxin and gibberellin synthesis were
3 Fig Tree Genome and Diversity 65

also differentially expressed. A study of FcFT1, a FLOWERING LOCUS T, was

previously performed by Ikegami et al. (2013b)⁠. They observed higher mRNA lev-
els in the ‘Houarishi’ variety leaves at the 1st to 6th nodes than higher nodes.
However, fruit-bearing and FcFT1-expressing nodes did not correspond completely,
as FcFT1 expression was detected even in basal nodes that usually bear no fruits in
figs. An additional flowering-related gene FLORICAULA/LEAFY (FcLFY), was
identified and characterized in the ‘Brunswick’ variety by Ma et al. (2020)⁠. The
highest expression was observed in apical buds, followed by flower buds and leaf
buds, which confirmed their involvement in flower bud differentiation and apical
meristem formation.

4.2.4 Response to Abiotic Factors

In addition to previously mentioned studies that tackle the plant and fruit develop-
mental biology, studies related to the physiological response to abiotic and biotic
factors represent an essential part of analyses (Kim et al., 2003; Kitajima et al.,
2018; Mascellani et al., 2021; Nawade et al. 2020a, b⁠; Sadder et al., 2021; Vangelisti
et al., 2019; Zhai et al., 2021).
In order to study the mechanism involved in response to salinity, Vangelisti et al.
(2019)⁠ analyzed transcriptomes from leaves of the variety ‘Dottato’ after irrigation
of the trees with 100 mM salinity concentration. As a result, already known salt-
regulated genes and some genes that have not been previously reported to be
involved in plant salinity stress were identified. The results can be used to develop
potential markers in plant breeding programs.
The effect of salinity stress on genes involved in the synthesis, accumulation, and
transport of soluble carbohydrates in ripe fruits of the variety ‘Dottato’ was investi-
gated by Mascellani et al. (2021),⁠ where a general increase in the transcript levels of
genes involved in the transport of soluble carbohydrates was observed.
Sadder et al. (2021)⁠ studied the differential expression of five putative salinity
responsive genes in three common fig landraces (‘Mwazi’, ‘Zraki’, and ‘Khdari’).
Based on the combined physiological and molecular data, FcSORD and FcDHN
genes were identified as potential molecular markers associated with stress toler-
ance response as they were able to distinguish tolerance levels.

4.2.5 Response to Biotic Factors

Plants pose different mechanisms to protect themselves against pests and diseases.
Latex, which is exuded from the wounded sites of the fig plant, contains several
toxic compounds. A cDNA for chitinase (CHI), a pathogenesis-related protein, was
isolated from the fig tree latex, and it was suggested that it could be involved in the
stress response caused by insect or pathogen attack (Kim et al., 2003)⁠. To infer
latex-associated defense mechanisms, a comparative study of transcriptomes
obtained from latex from immature fruit, young petioles, and lignified trunks was
66 D. Bandelj et al.

performed by Kitajima et al. (2018)⁠. Differentially expressed unigenes coding pro-

teins with known toxicity to pests and unigenes with no previously reported anti-
pest functions were identified, representing possible candidates for novel
defense-related proteins or chemicals.
It was observed previously that ficins are, in addition to 20% gum, 70% water,
and 1% other dry materials, a major component of fig latex (Zare et al., 2013)⁠.
Ficins are a class of papain-like cysteine proteases (PLCPs) which are the most
abundant family of cysteine proteases in plants, involved in abiotic and biotic stress
responses, growth, and senescence. Increased abundance in receptacles of
commercial-ripe fruit was observed, and genes encoding the main ficin isoforms
were characterized by Zhai et al. (2021)⁠. Ficin is used in the food and pharmaceuti-
cal industry, and to optimize the production of ficin, a gene was cloned from the
variety ‘Sabz,’ and recombinant protein was expressed in E. coli (Sattari et al., 2020)⁠.
Insights into fig-insect interactions (i.e., the black fig fly, Silba adipata) and
molecular mechanisms of terpene biosynthesis related to insect-plant interaction
were provided by Nawade et al. (2020a)⁠. In addition, Nawade et al. (2020b)⁠ studied
the emission of apocarotenoid volatiles and identified and characterized carotenoid
cleavage dioxygenase (FcCCD) genes involved in apocarotenoid biosynthesis.
They observed significant changes in apocarotenoids emitted over the development
process, which may have consequences for the interaction of fig-insect (including
fig wasp).

5 Conclusions

The genetic resources of figs consist of many varieties and ecotypes that arose spon-
taneously under distinct ecological pressures and through the human selection of
trees with attractive traits. The spontaneous hybridization of cultivated and wild
types and their further propagation also contributed to today's varieties' diversity.
Despite the research efforts in individual countries, a global catalog of figs has not
yet been prepared. Therefore, it is necessary to emphasize that comparison of fig
diversity from distinct countries is a difficult task, and the differences may be
reflected in: (1) different forms of fig under investigation (cultivated vs. wild), (2)
the number of analyzed samples, (3) selection of DNA markers and protocols, and
(4) history, practices, and traditions of fig cultivation. Therefore, establishing a con-
sortium of the fig research community with more target-oriented and coordinated
activities regarding the management of global common fig genetic resources is nec-
essary. Such collaborative organization would contribute to faster progress and
novel achievements in global fig genetic resources.
In general, fig genetic resources are rich, although different levels of genetic
diversity of cultivated figs and caprifigs have been reported in fig-producing coun-
tries. The genetic structure of figs has evolved differently over millennia of cultiva-
tion and reflects human activities, including selection and propagation processes,
3 Fig Tree Genome and Diversity 67

the pressure of ecological factors, and, ultimately, the introduction of varieties from
other countries in a defined region. As a result, genetic diversity studies do not nec-
essarily show a clear distinction or similarity of varieties according to geographical
origin. The lack of standardized protocol and a recommendation list of highly infor-
mative DNA markers appropriate for fig genetic resources evaluation also prevents
direct comparison of fig genetic diversity and the discovery of international syn-
onyms and homonyms.
SSRs were the most frequently used in fig diversity studies among all available
DNA markers. A review of the fig studies involving microsatellite markers has
shown that individual countries have used different combinations of microsatellites,
and in some studies, less informative loci were employed. In these cases, the choice
of loci certainly influenced the assessment of the degree of diversity of figs. The
latter suggests that a standardized list of the most informative microsatellite markers
for genetic studies of figs should be prepared. Uniform protocol and a standardized
list of microsatellites would better compare fig genetic resources among fig produc-
ing regions around the world and help address international synonyms of varieties.
The choice of the most appropriate markers is an important step in identifying vari-
eties. It affects the potential for discrimination of fig varieties, the rapidity and
extent of molecular analysis, and, consequently, the cost of the procedure.
Microsatellites are still the choice marker for variety identification, mainly due to
their high reproducibility and unambiguously readable allelic patterns. Among all
available microsatellites, choosing loci with the highest PIC and the lowest PI val-
ues from the published scientific reports is recommended.
Some studies have reported that the discrimination power of a defined DNA
marker system could be different concerning the genetic background of genotypes.
When closely related individuals such as clones are under investigation, finding dif-
ferences among them is higher with DNA markers that, in a single assay, cover a
larger part of the genome (for example, AFLPs). However, the complexity of inter-
pretation of banding patterns makes these dominant markers a less-preferred tool
for variety identification. With the increasing availability of fig genome and tran-
scriptome sequences, SNPs will undoubtedly contribute to identifying varieties and
genetically close-related genotypes by identifying novel alleles. A few studies also
reported that with the combination of different DNA marker systems, higher dis-
crimination power was obtained compared to a single DNA marker system used.
Despite the popularity of DNA markers in genetic studies, phenotyping with detailed
morphological description remains the essential and ‘must’ step in identifying and
characterization fig genetic resources.
The availability of high-throughput sequencing technologies has contributed to
the generation of new knowledge related to the biological process involved in plant
growth, stress response, development of different fig fruit types, buds differentia-
tion, and fruit ripening. A relatively small genome has also made the common fig
attractive as a model organism for studies focused on basic research. The high num-
ber of genomic and transcriptomic studies performed in recent years and the results
will contribute to the more efficient development of new varieties either with
68 D. Bandelj et al.

conventional breeding techniques or with techniques based on recombinant DNA

technology (e.g., CRISPR/Cas9) as first attempts have already been performed.

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Chapter 4
Genetic Diversity of Fig Varieties

Rim Ben Abdallah, Imed Othmani, Amel Lagha, and Sami Fattouch

Abbreviations

AMOVA Molecular Variance Analysis

ANOVA Analysis of variance
GO Genetic Ontology
ISSR Inter-Simple Sequence Repeats
Itraq Isobaric Tags for Relative and Absolute Quantitation
LTR Long Terminal Repeat
NCGR National Clonal Germplasm Repository
PCA Principal Component Analysis
PCR Polymerase Chain Reaction
RAPD Random Amplified Polymorphism DNA
RFLP Restriction Fragment Length Polymorphism
SAS Statistical Analysis System
SSR Simple-Sequence Repeats
UPGMA Unweighted Pair-Group Approach of Arithmetic Averages

R. Ben Abdallah (*) · I. Othmani · A. Lagha · S. Fattouch

EcoChem Laboratory, Department of Biological and Chemical Engineering, National
Institute of Applied Sciences and Technology (INSAT), University of Carthage (UCAR),
Tunis, Tunisia
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 77

M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_4
78 R. Ben Abdallah et al.

1 Introduction

The fig (Ficus carica L.) is one of the world’s oldest cultivated fruit species. The
English word fig comes from the Latin word ficus, which means “fig tree”. Caria, a
historic location of Asia Minor known for figs, is referenced in the species name
carica (Ferguson et al., 1990). The cultivation of figs was developed by the
Phoenicians, Carthaginians, and Greeks. The Romans spread it throughout the
Mediterranean basin (Roger, 2003). The fig tree progressively spread worldwide,
reaching far-flung places like China, India, Ethiopia, Arabia, Australia, and the
Americas, where it was introduced and widely propagated. The fig tree Ficus carica
L. developed from a monoecious species to a functionally dioecious species. Mgyz
(Persians), erineos (Greeks), teb (Egyptians), caprificus (Romans), and ettine or
dhokkar (Egyptians) are some of the names for the fig tree (Arabs). In Mediterranean
regions, the Arabs developed fig production. The Portuguese named them figo de
toco, while the Italians called profico or caprificus (Lahbib, 1984). Ficus carica L.,
a subtropical fruit tree, belongs to the genus Ficus, mostly found in the tropics
(Ferguson et al., 1990). Figs are grown in most warm and temperate regions and
have been for many years as a Mediterranean fruit tree (Sahin, 1997). Figs are com-
monly grown with other fruit trees, including olives, almonds, pomegranates, and
grapevines (Aljane, 2011). The common fig (Ficus carica L.) is a member of the
Moraceae family, which includes approximately 1400 species divided into 60 gen-
era. 2n 2x 26 is the number of somatic chromosomes (Ferguson et al., 1990). There
are about 700 species of Ficus in the genus Ficus (Berg, 2003). The fruits of this
species are drupelets that emerge from the ovaries in a closed syconium. The tiny fig
wasp pollinator (Blastophaga psenes L.) enters the fruit through a small opening
(ostiole). The only member of the genus Ficus farmed for its fruit is Ficus carica
(Ferguson et al., 1990). According to Smartt (1976), four types of figs are grown
worldwide: (a) The most familiar variety, which produces infertile seeds and pro-
duces figs without pollination. Some cultivars produce an excellent first crop (breba
crop) and then a good second crop (main crop); (b) the San Pedro type produces a
first crop (breba crop) that persists to ripening and is parthenocarpic. Pollination is
required to maturation the second crop (main crop). This variety‘s specialty is the
breba crop; (c) Smyrna type provides full fruit with viable seeds and requires pol-
lination with pollen from caprifigs (caprification). Though, the fruits will fall from
the tree before they mature if pollination is not done; (d) The Caprifig type produces
syconia flowers, which contain pollen that will pollinate the Smyrna in addition to
San Pedro varieties.
Several prospections and alternative methodologies for genetic resource man-
agement were studied to protect fig genetic resources. Pomological and morpho-
logical features, molecular markers, Biochemical traits, phytochemical contents
diversity, and proteomic and transcriptomic analysis were utilized to describe local
cultivars and assess genetic diversity. Various investigations on fig genetic resources
have recently been conducted around the regions. This chapter aims to present an
outline of fig agriculture, germplasm diversity, fig conservation, and usage improve-
ment using both traditional and modern approaches.
4 Genetic Diversity of Fig Varieties 79

2 Genetic Variability in Fig Germplasm

2.1 Morphological Variability

Morphological markers have typically been used to determine the amount and dis-
tribution of genetic diversity. Even though environmental factors and agronomic
methods substantially influence the expression of these markers, morphological
characterization is a highly suggested initial step to take before attempting more
in-depth biochemical or molecular studies. As a result, the morphological descrip-
tion remains the initial stage in agricultural plant genetic variety preservation.
Furthermore, plant inventories based on morphological descriptions provide the
first glimpse into the degree of phenotypic variation, which is critical for planning
genetic resource preservation strategies and establishing national collections
(Podgornik et al., 2010).
Botanical Description F. carica L. is a tree that grows to be 15–20 feet tall with
multiple spreading branches and a trunk that is more than 7 feet in diameter. The
plant’s latex is milky white and mostly made up of ficin, a protein hydrolytic
enzyme. Carica refers to the papaya-like leaves of the plant.

Leaf the leaves are bright green, single, alternating, and enormous, reaching a
length of up to one foot. They have a rough upper surface and a soft bottom and are
deeply lobed with 1–5 sinuses. Flowers are observed in containers; they emerge
from the axils of old leaves.

Flower male flowers occupy the lower part of the receptacle, while female flowers
occupy the upper section. The ripe receptacle, Saikonium, has a huge quantity of
little whitish seeds.

Seeds seeds range from 30 to 1600 per fruit and can be very small, small, medium,
or large. Unless fertilized, the seeds are edible and empty. The fertilized seeds give
the dried figs their distinctive nutty flavor. A white inner ring contains a seed mass
bonded with jelly-like flesh on the inside.

Fruit fruit figs are paired or solitary, axillary on leafy branchlets, and generally
pear-shaped. The fruit (fig) and reproductive systems of the Ficus species are mutu-
ally exclusive. Only their related agaonid wasps (Hymenoptera: Chalcoidea:
Agaonide) may pollinate it, and the wasps can only lay eggs in their linked fruit.
Pollinator wasps are required for successful pollination and reproduction of F. car-
ica species. On the other hand, Agaonid wasps require the presence of their linked
species of F. carica to reproduce successfully. In comparison to apple pulp, fruit
skin provided the majority of phytochemicals and antioxidant activity.
80 R. Ben Abdallah et al.

Root The plant’s root system is usually shallow and spread out. Roots can some-
times cover 50 feet of ground, although some roots can reach up to 20 feet in
porous soil.

Bark The bark is silky and smooth. The exterior bark is ash to silvery grey, with
irregular spherical flakes exfoliated. The center region of the bark is brownish or
light reddish-brown. Layers of pale yellowish or orange-brown colored granular
tissue make up the interior section (Gafoor et al., 2019).
The majority of descriptor lists for the characterization of genetic resources in
plants comprise many features whose evaluation is time-consuming and costly,
making the characterization of large germplasm collections challenging. As a result,
to make germplasm research and conservation easier, the most valuable factors for
each species must be carefully chosen. In addition, morphological characterization
is time-consuming; hence, descriptor lists for a wide range of species have been
created to help.
In order to make the characterization procedure in this species easier, a group of
international experts created and published a list of descriptors (International Plant
Genetic Resources Institute and Centre International de Hautes Etudes Agronomiques
Méditerranéennes, 2003). However, because of some unique characteristics of this
species, such as the frequent production of two different crops (breba and main
crop), the presence of four different types of leaves, and its complex floral biology,
morphological trait analyses in fig are even more complicated than in most fruit tree
species.
As a result, the current descriptors list has 192 properties, 126 of which are quali-
tative and 66 of which are quantitative. Because of the large number of features and
the requirement to replicate observations for at least two years, phenotypic charac-
terization in this species is time- and resource-intensive, particularly in developing
nations with limited resources.
Several studies have been carried out to determine the morphological and
pomological properties of cultivars of F. carica L. Darjazi (2011) reported physi-
cal characteristics and quality parameters of some local cultivars of fig (Ficus
carica L.) grown in Varamin, Iran. His results are presented in Table 4.1. At least
ten mature fruits and ten mature leaves from each of the fig cultivars were taken
from several areas of the same trees in Varamin Research Station during the last
week of July 2001. Darjazi (2011) conducted another study on the morphological
and pomological diversity of figs, in which 76 fig accessions were gathered in
Hatay, Turkey’s Eastern Mediterranean Region, in 2008 and 2009. Results are
shown in Table 4.2.
4

Table 4.1 Variation in morphological characteristics of fig cultivars in Varamin, Iran

Cultivar
Siah bolbol Hallavi
Traits Bidaneh Paizeh Zard riz Siah zoodras Siah diras Morabaii Hallavi riz dourosht
Leaf trait
Leaf length 18.75 20.503. 17.00 12.00 25.20 26.50 25.00 24.00 21.50
(cm)
Leaf width 12.60 16.18 13.75 8.25 14.80 17.00 18.60 16.20 14.60
(cm)
Leaf area 135 187 85 40 185 220 280 225 165
(LxW) (cm2)
Genetic Diversity of Fig Varieties

Number of 3 3 5 3 3 5 3 3 3
lobes
Length of 8.5 9.8 9.5 5.5 11.3 12.50 13 11.9 10.51
central lobe
(cm)
Leaf margin Regular Regular Irregular Regular Irregular Irregular Regular Irregular Irregular
toothed toothed toothed toothed toothed toothed toothed toothed toothed
Leaf color Light green Dark green Dark green Dark green Dark green Dark green Light green Dark green Dark green
Petiole length 4.75 5.50 3.75 4.10 7.00 12.00 6.00 4.75 4.75
(cm)
Petiole 2.60 3.60 2.75 3.25 3.75 3.50 3.40 3.60 2.80
thickness
(mm)
Depth of 2.40 3.20 3.75 2.25 3.50 4.70 4.40 2.80 2.80
upper sinuses
(cm)
Leaf trace 4.25 4.70 3.90 3.45 4.40 4.25 5.30 4.10 3.10
(mm)
(continued)
81
82

Table 4.1 (continued)

Cultivar
Siah bolbol Hallavi
Traits Bidaneh Paizeh Zard riz Siah zoodras Siah diras Morabaii Hallavi riz dourosht
Midvein 14 15 13 7.50 18.50 17.00 19.00 19.00 17.00
length (cm)
Mid lobe base 4.00 5.00 2.50 1.40 4.20 3.20 4.75 4.90 4.50
width (cm)
Angle of 110 76 90 110 0 0 85 0 0
central lobe
tip
Angle of 102 64 70 100 0 0 76 0 0
lateral lobe tip
Length of 10.50 12.30 10.50 6.00 12.70 12.70 14.60 13.00 11.80
lateral lobe
(cm)
Leaf blade Soft Soft Medium Soft Rough Rough Rough Medium Medium
surface
Leaf blade Fragile Flexible Flexible Fragile Fragile Fragile Fragile Fragile Fragile
texture
Upper sinuses U U V U V U U V V
shape
Density of None Intermediate None Intermediate None None None None None
hairs on leaf
upper surface
Bud trait
R. Ben Abdallah et al.
4

Length of 19.50 12.00 7.40 6.25 9.50 9.50 11.00 18.75 5.50
current season
growth (cm)
Diameter of 8.25 7.75 8.00 7.50 9.50 9.50 8.50 8.75 6.25
current season
growth (mm)
Terminal bud 9.50 11.90 10.30 6.40 10.80 9.50 12.50 8.30 8.50
length (mm)
Terminal bud 3.60 4.30 3.90 3.80 5.40 5.00 5.00 4.30 3.70
width (mm)
Tbl / Tbw 2.63 2.76 2.64 1.68 2.00 1.90 2.50 1.93 2.29
Genetic Diversity of Fig Varieties

Shoot 48 27 15 12 23 22 23 37 15
internode
length (mm)
Internode 8 6 7 8 8 8 7 9 5
number
Tree traits
Tree growth Weeping Erect Spreading Spreading Weeping Erect Weeping Weeping Weeping
habit
Tree vigor Medium Medium Low Low Medium Low High Medium Medium
Darjazi (2011)
83
Table 4.2. Variation in morphological characteristics of fig cultivars from the Eastern Mediterranean Region of Turkey
Number Number Length
Tree Shoot of leaves of fruits Leaf Leaf of center Leaf Petiole Petiole Number
growth length on a on a length width lobe area length thickness Leaf of leaf Apical Full Harvesting
Accession habit Tree vigor (cm) shoot shoot (cm) (cm) (cm) (cm2) (cm) (mm) shape lobes dominancy maturity period
Mor 1 Open High 40.0 11.2 7.2 22.8 21.8 13.3 320.1 8.5 5.7 D 5 Present 1–30 sept. 21–40
Mor 2 Open Intermediate 23.1 7.3 4.4 22.4 18.2 13.8 282.2 7.8 5.3 A 5 Present 1–30 sept. 41–60
Mor 3 Open Intermediate 11.8 6.3 5.4 20.5 17.1 12.6 256.4 6.2 4.3 C 5 Absent 11–31 Aug. 21–40
Mor 4 Weeping Intermediate 29.4 7.4 6.2 20.1 15.8 10.1 246.6 8.2 6.0 G 3 Present 1–30 sept. >60
Mor 5 Semierect Intermediate 15.9 6.1 3.5 19.8 17.2 11.9 259.8 5.4 6.1 D 5 Absent 1–10 Aug. 21–40
Mor 6 Open High 8.8 5.2 2.7 18.3 16.6 10.9 223.6 5.8 4.8 D 5 Absent 1–10 Aug. 21–40
Sultani 1 Open Intermediate 7.8 4.4 2.4 18.9 14.8 12.4 204.5 7.1 4.7 B 5 Absent 11–31 Aug. 21–40
Sultani 2 Open Intermediate 9.4 4.7 2.5 16.4 15.0 11.2 163.4 6.0 3.6 C 5 Absent 11–31 Aug. 21–40
Sultani 3 Open Intermediate 10.2 5.7 4.1 17.9 15.7 11.5 213.0 7.1 4.3 D 5 Present 11–31 Aug. 21–40
Kabak 1 Open High 11.5 6.3 4.7 26.6 22.1 15.3 348.6 8.5 5.9 C 5 Absent 1–10 Aug. 41–60
Kabak 2 Spreading Intermediate 10.6 6.5 4.8 23.4 21.2 12.8 325.8 7.3 5.6 C 5 Absent 1–10 Aug. 41–60
Şeble 1 Spreading High 26.1 8.8 5.7 20.6 18.8 12.8 278.5 8.1 4.6 C 5 Present 11–31 Aug. 21–40
Şeble 2 Open Intermediate 26.4 9.1 5.3 21.5 17.5 12.2 269.4 6.9 5.4 C 5 Present 11–31 Aug. 21–40
Kıreni 1 Open Intermediate 15.2 10.5 2.6 21.0 19.1 16.1 263.4 8.0 5.9 F 5 Absent 11–31 Aug. 21–40
Kıreni 2 Spreading Intermediate 13.6 6.5 4.5 21.4 18.0 12.6 280.4 9.1 5.8 B 5 Absent 1–10 Aug. 41–60
Sehli 1 Open Intermediate 9.3 5.2 3.4 27.6 19.5 13.0 276.3 7.4 5.1 C 5 Absent 1–10 Aug. 41–60
Sehli 2 Open Intermediate 9.0 7.5 2.6 17.1 14.6 11.0 176.6 5.9 5.0 D 5 Absent 1–10 Aug. 21–40
Meryemi 1 Open Low 18.0 7.1 4.9 19.9 17.9 12.2 221.7 7.5 4.4 B 5 Absent 1–10 Aug. 21–40
Meryemi 2 Spreading Intermediate 11.9 6.1 3.7 23.4 17.5 14.0 269.6 6.6 5.2 G 3 Absent 11–31 Aug. 21–40
Kuruye 1 Semierect Intermediate 11.9 4.9 4.2 20.2 17.5 12.4 235.8 6.9 5.1 D 5 Present 11–31 Aug. 41–60
Kuruye 2 Semierect Intermediate 12.7 7.3 5.2 18.8 16.5 12.1 219.7 5.9 4.8 D 5 Present 11–31 Aug. 41–60
Kırmızı 1 Semierect Intermediate 31.8 9.7 7.2 21.1 18.8 13.5 264.0 7.5 4.7 C 5 Present 11–31 Aug. 21–40
Kırmızı 2 Erect Low 8.1 5.1 2.7 16.5 13.7 10.3 172.5 5.2 4.6 G 3 Absent 1–10 Aug. 15–20
Lopkara 1 Open High 10.9 6.3 5.4 22.8 20.1 13.6 302.6 7.5 5.3 D 5 Absent 11–31 Aug. 21–40
Lopkara 2 Semierect Intermediate 33.3 7.5 5.2 24.1 21.1 13.2 346.2 9.6 5.5 G 3 Present 11–31 Aug. 21–40
Ramlı 1 Open High 10.7 5.4 2.7 16.6 14.6 10.0 166.0 4.7 4.1 C 5 Absent 1–10 Aug. 15–20
Ramlı 2 Semierect Low 9.7 5.4 3.3 17.2 14.1 10.5 166.8 4.8 4.2 G 3 Absent 1–10 Aug. 15–20
Bığrasi 1 Open High 20.9 7.2 5.3 24.3 22.6 13.2 358.8 7.3 5.3 G 3 Absent 11–31 Aug. 41–60
Bakrasi 2 Open Intermediate 15.2 4.2 4.2 17.5 17.4 9.8 205.3 5.0 4.3 G 3 Absent 11–31 Aug. 21–40
Bakras 3 Spreading Low 8.5 7.2 5.9 22.5 20.1 13.1 283.8 7.5 5.2 G 3 Present 11–31 Aug. 41–60
Bakrasi 4 Weeping High 18.6 7.4 6.1 21.0 21.1 11.0 306.5 7.1 4.5 G 3 Present 11–31 Aug. 41–60
Bakrasi 5 Semierect Intermediate 25.7 9.4 8.1 23.6 22.5 11.8 371.6 8.9 5.0 G 3 Present 11–31 Aug. 40–60
Şami Open Intermediate 24.5 10.4 7.5 23.5 20.7 12.8 280.5 8.2 5.4 C 5 Present 11–31 Aug. 41–60
Fransavi Open Low 20.2 6.0 3.7 20.3 17.1 11.4 250.3 8.4 4.8 G 3 Present 11–31 Aug. 41–60
Hılvıni Weeping High 15.9 5.8 3.4 19.3 16.5 13.4 219.4 5.1 4.6 A 5 Absent 11–31 Aug. 21–40
Büyük Siyahlop Semierect Intermediate 18.5 7.2 3.7 21.3 17.9 13.1 262.7 7.0 5.4 C 5 Present 11–31 Aug. 41–6
Sıhle Open High 9.1 6.3 3.9 20.4 17.5 13.3 243.7 5.3 4.8 C 5 Absent 11–31 Aug. 41–60
Kilis Erect Low 16.8 9.5 4.7 22.8 21.6 12.5 334.1 7.0 5.0 C 5 Present 1–10 Aug. 21–40
Ahmediye Weeping High 11.3 6.9 5.6 23.5 21.2 14.5 332.4 6.1 5.0 G 3 Absent 11–31 Aug. 41–60
Burnu Kızıl Semierect Low 9.0 5.4 4.3 19.9 17.4 11.0 283.2 6.0 4.5 G 3 Absent 11–31 Aug. 41–60
Allene Karası Semierect Intermediate 11.1 5.8 3.4 23.1 18.9 15.1 280.3 6.7 6.6 D 5 Present 1–10 Aug. 21–40
Beyaz Fahli Open Intermediate 22.2 7.5 3.9 21.6 16.7 13.6 253.5 6.0 5.4 A 5 Present 11–31 Aug. 21–40
Kandamık Spreading Low 9.4 6.5 3.3 21.5 18.0 12.4 316.1 5.3 5.6 C 5 Present 1–10 Aug. 21–40
Fahli Semierect Intermediate 10.9 5.8 2.6 19.0 16.0 12.5 221.2 6.1 5.6 C 5 Present 11–31 Aug. 21–40
Fetike Open Intermediate 9.8 6.2 3.0 19.4 16.2 12.3 246.1 7.3 5.2 B 5 Present 11–31 Aug. 40–60
Armut Sapı Open Low 6.5 5.8 2.8 17.2 16.1 10.9 195.7 6.5 4.5 D 5 Absent 1–10 Aug. 21–40
Payas Spreading Intermediate 8.4 4.8 2.6 19.6 15.9 12.0 223.8 6.8 3.7 B 5 Present 11–31 Aug. 41–60
Gud Yeniği Semierect Low 9.2 6.4 4.3 21.1 17.0 14.1 237.3 6.4 4.8 D 5 Present 1–10 Aug. 21–40
Baldır Open Intermediate 9.4 5.7 4.1 20.1 15.1 11.5 207.0 9.8 4.1 G 3 Absent 11–31 Aug. 41–60
Halep Semierect Intermediate 25.8 6.6 4.5 25.1 23.5 13.5 347.0 11.4 6.4 G 3 Present 11–31 Aug. 41–60
Erkenci Semierect Intermediate 8.9 6.6 4.3 20.7 16.2 14.6 223.6 5.8 4.1 C 5 Absent End of July 21–40
(continued)
Table 4.2 (continued)
Number Number Length
Tree Shoot of leaves of fruits Leaf Leaf of center Leaf Petiole Petiole Number
growth length on a on a length width lobe area length thickness Leaf of leaf Apical Full Harvesting
Accession habit Tree vigor (cm) shoot shoot (cm) (cm) (cm) (cm2) (cm) (mm) shape lobes dominancy maturity period
Yeşil İncir Open Intermediate 32.1 9.2 5.3 21.6 18.5 12.6 303.2 5.3 5.4 A 7 Present 11–31 Aug. 21–40
Şebli Spreading Intermediate 21.8 5.6 3.3 19.9 17.9 11.8 271.2 6.1 4.7 G 3 Absent 11–31 Aug. 21–40
Tınesvit Weeping Intermediate 15.0 5.8 4.3 21.7 18.4 13.0 254.9 6.6 4.7 C 5 Absent 11–31 Aug. 21–40
Sütlü Sarı Open Intermediate 18.0 7.5 4.2 22.8 19.5 12.4 281.5 6.7 5.1 G 3 Present 11–31 Aug. 21–40
Mersinli Open Intermediate 9.6 5.1 3.0 23.0 16.6 14.3 231.6 6.9 5.4 C 5 Present 1–10 Aug 21–40
Zırhıni Weeping High 19.1 7.2 5.1 22.4 19.0 13.3 259.5 8.6 5.1 A 5 Absent 11–31 Aug. 21–40
Şibili Open Intermediate 15.7 8.5 6.6 21.6 19.9 12.2 311.8 7.1 5.7 F 5 Absent 11–31 Aug. >60
Karagöz Erect Intermediate 25.8 8.1 5.4 20.8 18.4 10.7 273.9 10.2 4.8 G 3 Present 11–31 Aug. 41–60
Beyaz İncir Open Intermediate 14.2 6.3 4.2 21.8 17.3 13.5 280.1 7.4 4.8 D 5 Absent 11–31 Aug. 21–40
Sarı 1 Spreading Intermediate 20.9 8.2 6.0 23.9 21.0 13.5 336.0 8.1 6.0 C 5 Present 11–31 Aug. 21–40
Sarı 2 Open Intermediate 16.6 6.9 2.8 19.3 16.5 10.9 287.3 7.4 5.4 D 5 Present 1–10 Aug. 21–40
Sarı 3 Open Low 7.9 6.8 4.3 20.9 19.0 13.5 259.3 10.1 4.1 B 5 Absent 1–10 Aug. 21–40
Sarı 4 Semierect High 19.4 10.4 7.2 22.8 21.6 11.9 356.9 9.7 5.0 G 3 Present 11–31 Aug. 41–60
Sarı 5 Open Intermediate 33.3 10.3 6.4 21.9 18.3 12.2 249.1 5.0 5.0 A 7 Present 11–31 Aug. 21–40
Sarı 6 Open High 15.4 7.1 4.7 21.2 19.2 12.4 288.1 8.2 5.3 G 3 Absent 1–30 sept. 21–40
Siyah 1 Open Low 15.4 6.4 3.8 23.5 20.8 13.5 296.7 8.4 5.4 G 3 Absent 11–31 Aug. 21–40
Siyah 2 Semierect Low 7.5 4.9 3.1 21.1 17.9 14.0 277.6 6.0 5.6 D 5 Present 11–31 Aug. 21–40
Siyah 3 Weeping Intermediate 17.8 6.6 5.0 21.8 18.0 14.0 280.4 8.0 4.6 C 5 Present 11–31 Aug. 21–40
Siyah 4 Spreading Intermediate 19.8 7.0 5.4 22.2 18.6 12.7 301.0 8.5 5.7 C 5 Absent 1–30 sept. 41–60
Siyah 5 Open Low 14.9 5.7 3.6 21.6 19.8 12.3 275.5 7.1 4.4 G 3 Absent 11–31 Aug. 21–40
Siyah 6 Weeping High 22.1 5.0 3.7 19.0 17.2 11.6 221.6 6.7 4.6 C 5 Absent 11–31 Aug. 21–40
Siyah 7 Weeping High 19.3 7.4 5.2 23.1 18.4 13.8 273.3 11.0 5.2 A 5 Present 11–31 Aug. 21–40
Siyah 8 Spreading High 32.1 6.4 3.6 20.7 18.1 11.1 262.3 10.4 4.8 G 3 Present 1–30 sept. >60
Çalişkan and Atila (2012)
4 Genetic Diversity of Fig Varieties 87

2.2 Pomological Traits

Darjazi (2011) has reported pomological properties and quality parameters of some
local cultivars of fig (Ficus carica L.) grown in Varamin, Iran. Fruits were evaluated
for pomological purposes using a set of criteria with physical attributes and qualita-
tive factors. Three replications are considered; in each replication, there are 10
fruits. Results are reported in Table 4.3 in a study of the morphological and pomo-
logical variety of figs in Hatay, Turkey’s Eastern Mediterranean Region, in which 76
fig accessions were gathered in 2008 and 2009. Results are reported in Table 4.4.

2.3 Agronomical Parameters

Different agronomical parameters such as annual yield, cumulative yield, yield effi-
ciency, ripening period, harvesting of brebas and figs, and trunk cross-sectional area
were studied (Pereira et al., 2015, 2017) to investigate the agronomic behavior and
quality of different fig cultivars produced in Spain for fresh consumption. The
researched fig cultivars were ‘Cuello Dama Blanco’, ‘Brown Turkey’, ‘Colar Elche’,
‘SanAntonio’, ‘Banane’, and ‘Blanca Bétera’.
Results showed that the ripening period for brebas was 9–21 days, while the
main crop was 67–75 days. The earliest maturing cultivar was ‘San Antonio,’ which
reached maturity in the first week of June. ‘Brown Turkey’ and ‘Blanca Bétera’
breba fruit ripened in mid-June, followed by ‘Banane’, ‘Cuello Dama Blanco’, and
‘Colar Elche’ breba fruit in the third week of June. San Antonio, Brown Turkey, and
Blanca Bétera’s second crops matured in mid-July, while ‘Banane,’ ‘Cuello Dama
Blanco,’ and ‘Colar Elche’ matured at the end of July. The maturation period for
these six cultivars lasted until early October, when temperatures decreased and the
first autumn rains began. Each cultivar and block’s annual breba crop production
(kg/tree) was much lower than the main crop‘s yearly yield, with unpredictable
values between years and blocks. The cultivars ‘San Antonio’ and ‘Brown Turkey’
had the largest annual breba crop yield, with maximum values of 5.5 and 5.4 kg/tree
on the fifth green, respectively (2012). Annual primary crop production improved
year after year, reaching the sixth green (2013) with values of 83.2, 74.6, 60.2, 56.7,
53.7, 27.2 kg/tree for ‘Banane’, ‘BrownTurkey’, ‘Colar Elche’, ‘Cuello Dama
Blanco’, ‘San Antonio’, and ‘Blanca Bétera’. Furthermore, a high percentage of
damaged fruit was discovered during the harvest of brebas and main crop figs, rang-
ing from 30% to 11% depending on the brebas and main crop figs; the dark cultivars
had the highest percentage of damaged fruit.
The annual yield also differed between blocks, which delayed the trees’ entry
into production due to harsh pruning throughout the winter. During the study period
of four years (2010–2013), the cumulative yield of the breba crop (kg/tree) was
considerably lower than the cumulative yield of main crop figs during the same
period. The cultivars ‘San Antonio,’ ‘Banane,’ and ‘Brown Turkey’ yielded the most
brebas, with 17.4, 17.2, and 10.8 kg/tree, respectively, while the cultivar ‘Cuello
Table 4.3 Variation in pomological characteristics of fig cultivars in Varamin, Iran
88

Cultivar
Siah bolbol Hallavi
Traits Bidaneh Paizeh Zard riz Siah zoodras Siah diras Morabaii Hallavi riz dourosht
Fruit trait
Fresh fruit 27.9 43.5 24.4 8 33.5 30.7 8.6 19.7 20.7
weight (g)
Dried fruit 5 7.4 4.1 1.6 4.9 4.9 1.6 3.4 4.4
weight (g)
Ffw /Dfw 0.17 0.16 0.16 0.19 0.14 0.15 0.18 0.16 0.21
Fruit volume 31.2 46.6 24.3 7.4 33.7 31.4 9.8 19.7 23.7
(cm3)
Fruit diameter 38 45 35 21 40 38 24 32 37
(mm)
Fruit length 27 35 23 20 31 25 25 36 22
(mm)
Fruit shape 1.40 1.52 1.28 1.05 1.29 1.52 0.98 0.88 1.68
index(Fd/Fl)
Fruit shape Oblate Oblate Oblate Globose Oblate Oblate Globose Oblong Oblate
Ostiole width 2.9 2.7 4.2 1.6 4.6 4.2 0 3.1 4
(mm)
Stalk length 7.00 6.00 11.5 15 4.00 5.00 5.80 11.0 12.0
(mm)
Stalk 5.20 5.30 4.00 1.70 4.80 4.50 3.80 4.30 4.60
diameter
(mm)
Sugar (%) 15.3 9.8 16.2 16.5 15 16.8 16.5 18.9 17.9
Total soluble 18 13.3 18.4 28.5 20 22.5 28.2 24 17.1
solid (%)
R. Ben Abdallah et al.
4

Fruit skin Light-green Canary Yellow-green Purplish Purple Black Canary Light-brown Light-brown
over color yellow brown yellow
Fruit ribs None None None None None None None Prominent Prominent
Bloom Medium None None None None None None None None
Fruit skin None None Minute None None None None None None
cracks
Ease of Easy Easy Easy Easy Difficult Easy Difficult Easy Easy
peeling
Fruit flesh Light yellow Light yellow Light yellow White White White Light yellow Light yellow Light yellow
color
Pulp internal Amber Light Dark Amber Amber Amber Amber Amber Amber
Genetic Diversity of Fig Varieties

color strawberry strawberry

red red
Fruit cavity Medium Medium Small Small Small Small None Small Medium
Pulp juiciness Juicy Little juicy Little juicy Little juicy Little juicy Little juicy Little juicy Little juicy Little juicy
Shape of the Short thick Short thick Long slender Long slender Short thick Short thick Short thick Long slender Short thick
fruit stalk
Pulp flavor Aromatic None None None None None None None None
Seed number Low Medium High Low Low Medium Low Low Medium
Neck length None None None None None None Short Short None
Scale color of Light-orange Purplish pink Red with Light Dark- purple Dark- purple Canary Light-brown Light-brown
eye white margins yellow- white with white yellow
margins
Stalk color Green Yellow-green Yellow-green Light-yellow Dark-green Dark-green Light-yellow Dark-green Dark-green
Number of 1 1 1 1 1 1 2 1 1
fruit per node
Breba crop High Low Meduim None None None High None None
Skin Medium Medium Medium Thin Medium Thin Thin Thin Thin
thickness
89

Darjazi (2011)
Table 4.4 Variation in pomological characteristics of fig cultivars from the Eastern Mediterranean Region of Turkey
90

Fruit Fruit Neck Ostiole Fruit skin Total

Fruit diameter length length width thickness soluble TSS/ Fruit
Accession weight (g) (mm) (mm) Index (mm) (mm) (mm) solids (%) Acidity shape Skin color Pulp color
Şami 53.1 49.1 56.3 0.9 9.0 4.1 1.8 20.6 176.3 Globose Green Amber
Fransavi 48.9 46.1 46.4 1.0 5.3 4.0 1.4 19.8 77.1 Globose Green Red
Hılvıni 21.6 33.3 32.4 1.0 3.4 1.3 1.2 22.7 163.9 Globose Green Amber
Büyük 40.4 41.3 42.8 1.0 4.6 5.1 1.3 20.6 141.4 Globose Purple Amber
Siyahlop
Sıhle 38.2 40.6 45.7 0.9 7.2 1.9 1.4 22.5 117.9 Globose Brown Amber
Kilis 71.6 55.5 47.1 1.2 4.1 3.5 1.1 20.2 81.1 Oblate Green Amber
Ahmediye 78.2 52.6 52.3 1.0 4.7 1.3 1.6 20.8 115.3 Globose Green Pink
Burnu Kızıl 52.5 46.2 44.5 1.0 3.0 8.1 1.2 19.0 68.5 Globose Green Red
Allene Karası 45.6 47.0 38.7 1.2 4.0 6.2 1.3 19.2 129.6 Oblate Black Amber
Beyaz Fahli 40.7 47.2 36.2 1.3 3.5 21.0 1.1 23.7 127.5 Oblate Green Pink
Kandamık 58.1 49.2 39.4 1.2 1.9 1.9 1.1 20.0 94.2 Oblate Green Pink
Fahli 37.0 42.1 34.9 1.2 3.9 6.0 1.4 23.4 127.8 Oblate Green Pink
Fetike 59.4 49.0 42.9 1.1 5.7 8.3 1.6 20.5 156.3 Globose Green Pink
Armut Sapı 34.0 40.0 41.1 1.0 6.0 4.6 1.5 20.0 87.6 Globose Yellow Pink
Payas 18.9 32.5 31.0 1.0 0.5 1.9 1.2 20.4 108.4 Globose Green Dark red
Gud Yeniği 32.7 40.2 29.8 1.4 0.0 5.4 1.0 22.5 131.5 Oblate Green Amber
Baldır 40.5 42.3 39.6 1.1 5.6 3.2 1.6 20.5 81.1 Globose Green Red
Halep 56.2 50.3 44.6 1.1 8.1 4.5 1.6 18.4 110.9 Globose Green Amber
Erkenci 39.6 41.8 44.8 0.9 5.7 4.9 1.6 20.9 150.0 Globose Brown Amber
Yeşil İncir 66.5 51.8 43.0 1.2 2.3 3.3 1.9 22.5 70.9 Oblate Green Dark red
Şebli 38.6 39.1 42.3 0.9 2.3 3.7 1.0 23.5 155.1 Globose Brown Amber
Tınesvit 37.1 38.4 51.4 0.7 8.5 1.5 1.5 22.0 119.5 Oblong Brown Amber
Sütlü Sarı 51.1 47.0 50.2 0.9 7.5 2.4 1.4 20.4 168.6 Globose Yellow Pink
R. Ben Abdallah et al.

Mersinli 56.6 48.5 44.6 1.1 2.9 4.9 1.4 20.7 123.9 Globose Yellow Red
4

Zırhıni 22.9 34.7 46.6 0.7 7.1 1.3 1.1 22.4 118.2 Oblong Brown Red
Şibili 42.9 41.7 49.5 0.8 13.0 4.7 1.1 20.2 172.8 Oblong Purple Pink
Karagöz 46.6 42.9 47.4 0.9 5.8 1.1 1.2 20.2 69.5 Globose Purple Dark red
Beyaz İncir 46.8 45.7 41.7 1.1 5.1 5.8 1.4 20.4 79.0 Globose Green Dark red
Sarı 1 81.6 56.7 50.3 1.1 9.5 4.5 2.3 18.4 75.9 Globose Yellow Dark red
Sarı 2 62.3 50.1 52.8 0.9 7.8 6.7 1.8 19.7 199.8 Globose Yellow Pink
Sarı 3 54.3 48.6 43.1 1.1 8.3 6.8 1.7 20.8 104.1 Globose Yellow Pink
Sarı 4 37.5 38.0 40.3 0.9 4.0 5.9 1.9 17.5 82.0 Globose Green Pink
Sarı 5 74.0 53.8 44.1 1.2 3.1 3.0 2.0 21.4 80.7 Oblate Yellow Red
Sarı 6 87.4 58.7 48.4 1.2 4.3 5.1 1.4 23.6 132.0 Oblate Yellow Amber
Genetic Diversity of Fig Varieties

Siyah 1 58.4 50.2 47.6 1.1 5.5 2.5 1.7 19.9 107.9 Globose Brown Red
Siyah 2 36.0 42.5 36.5 1.2 2.7 3.8 1.0 21.9 102.4 Oblate Black Pink
Siyah 3 24.8 36.1 31.3 1.2 0.5 3.2 1.0 19.7 59.5 Oblate Black Dark red
Siyah 4 80.4 55.4 47.5 1.2 3.7 6.0 1.5 18.3 55.1 Oblate Black Red
Siyah 5 21.3 32.6 36.1 0.9 2.8 0.6 0.6 22.3 117.7 Globose Black Pink
Siyah 6 29.0 38.0 44.6 0.9 6.4 2.2 1.4 22.4 156.2 Globose Brown White
Siyah 7 35.6 38.8 47.8 0.8 7.8 2.4 1.8 21.0 91.5 Oblong Brown Pink
Siyah 8 40.1 40.6 44.3 0.9 5.0 1.1 1.2 21.5 67.2 Globose Purple Dark red
Mor 1 66.8 52.2 50.7 1.0 8.2 3.6 2.1 20.7 100.0 Globose Brown Red
Mor 2 65.3 49.9 48.4 1.0 3.3 4.6 1.8 19.4 104.8 Globose Purple Pink
Mor 3 36.2 39.5 47.4 0.8 7.4 2.0 1.7 20.8 147.1 Oblong Brown Amber
Mor 4 46.6 44.1 43.7 1.0 4.6 1.9 1.1 21.8 64.3 Globose Purple Red
Mor 5 30.9 37.0 38.3 1.0 2.7 3.6 1.3 22.6 204.3 Globose Purple Amber
Mor 6 45.3 45.7 36.1 1.3 2.5 2.8 1.4 18.5 108.1 Oblate Brown Red
Sultani 1 35.3 40.1 40.4 1.0 4.2 3.2 1.9 25.5 170.5 Globose Yellow Pink
Sultani 2 50.4 46.6 46.7 1.0 6.0 5.0 2.2 23.0 114.7 Globose Yellow Red
Sultani 3 44.1 45.8 43.7 1.0 6.3 3.5 1.3 19.2 81.9 Globose Green Red
91

(continued)
92

Table 4.4 (continued)

Fruit Fruit Neck Ostiole Fruit skin Total
Fruit diameter length length width thickness soluble TSS/ Fruit
Accession weight (g) (mm) (mm) Index (mm) (mm) (mm) solids (%) Acidity shape Skin color Pulp color
Kabak 1 81.7 55.8 54.5 1.0 8.0 2.2 1.7 17.9 51.4 Globose Green Dark red
Kabak 2 99.4 61.1 46.4 1.3 7.5 4.6 2.2 19.0 81.2 Oblate Green Dark red
Şeble 1 51.9 43.7 50.8 0.9 3.9 5.0 1.7 19.1 105.9 Globose Brown Pink
Şeble 2 50.7 44.5 46.8 1.0 4.7 6.7 1.2 19.4 99.5 Globose Brown Pink
Kıreni 1 47.7 46.9 43.4 1.1 3.9 10.3 1.8 21.2 112.1 Globose Yellow Pink
Kıreni 2 69.7 51.8 52.4 1.0 6.4 4.4 1.3 22.0 195.4 Globose Yellow Pink
Sehli 1 33.7 38.1 43.0 0.9 5.5 2.1 1.4 23.4 107.3 Globose Brown Amber
Sehli 2 23.1 32.2 38.9 0.8 4.8 2.1 1.2 24.3 144.0 Oblong Brown Amber
Meryemi 1 55.3 49.2 50.1 1.0 8.7 6.8 1.8 19.2 94.4 Globose Yellow Dark red
Meryemi 2 47.6 45.5 39.4 1.2 3.1 5.1 1.2 20.6 77.7 Oblate Green Pink
Kuruye 1 25.2 34.5 42.2 0.8 7.0 2.6 1.5 23.0 145.8 Oblong Brown Amber
Kuruye 2 21.2 30.9 43.4 0.7 8.7 2.0 1.4 24.5 230.6 Oblong Brown Amber
Kırmızı 1 28.6 36.9 46.9 0.8 7.9 1.3 1.6 22.4 149.6 Oblong Brown Red
Kırmızı 2 12.3 27.7 28.4 1.0 1.7 2.8 1.5 27.1 136.8 Globose Brown Red
Lopkara 1 30.5 35.4 42.7 0.8 6.8 1.4 1.9 22.4 161.1 Oblong Brown Pink
Lopkara 2 54.4 46.6 47.1 1.0 5.8 6.3 1.7 19.5 69.5 Globose Black Red
Ramlı 1 24.6 33.3 37.9 0.9 5.4 4.9 1.1 20.0 145.4 Globose Green Pink
Ramlı 2 18.0 29.7 34.6 0.9 2.3 4.4 0.9 22.3 127.8 Globose Green Pink
Bığrasi 1 29.3 37.0 37.3 1.0 2.0 3.4 1.0 21.2 74.2 Globose Green Pink
Bakrasi 2 22.4 37.5 36.6 1.0 1.3 4.6 1.2 19.7 90.2 Globose Green Red
Bakras 3 34.0 38.8 39.7 1.0 2.8 2.8 1.2 20.4 82.0 Globose Green Pink
Bakrasi 4 45.0 43.0 47.0 0.9 4.5 4.4 1.8 16.0 63.6 Globose Green Pink
Bakrasi 5 39.3 41.8 49.8 0.8 3.4 5.0 1.7 19.5 100.9 Oblong Green Pink
Çalişkan and Atila (2012)
R. Ben Abdallah et al.
4 Genetic Diversity of Fig Varieties 93

Dama Blanco’ yielded the least, with 4.2 kg/tree. The cultivars ‘Banane’ and ‘Brown
Turkey’ had the largest cumulative yields of main crop figs, with
229 and 186 kg/tree, respectively. The cultivar ‘Blanca Bétera’, on the other
hand, had the lowest cumulative yield of 65.8 kg/tree.
During the sixth green, the trunk cross-sectional area connected to tree vigour
differed amongst cultivars. With a trunk section of 93.3 cm2, the cultivar ‘Cuello
Dama Blanco’ was the most vigorous, followed by ‘Banane’ and ‘Colar Elche’ with
90.5 and 79 cm2, respectively. According to the International Plant Genetic
Resources Institute (IPGRI), these three cultivars have only moderate vigour under
these growing conditions. Regardless, these varieties are regarded as prolific culti-
vars. The ‘Colar Elche’ cultivar, on the other hand, has a spreading growth tendency.
Hence commercial plantations for this cultivar would need to have larger tree spac-
ing, such as 6 m x 6 m, to minimize tree overlap.
Yield efficiency (kg/cm2), calculated as the ratio between the cumulative yield
and the trunk cross-sectional area, differed significantly among cultivars. Brebas
presented values lower than main crop figs due to the low annual yields. The ‘Brown
Turkey’ and ‘Banane’ cultivars exhibited good yield efficiency with 2.7 and 2.6 kg/
cm2, respectively, followed by the ‘San Antonio’ and ‘Colar Elche’ cultivars with
1.9 and 1.6 kg/cm2, respectively.

2.4 Molecular and Biochemical Variability Assessment

2.4.1 Structure and Composition of Fig Genome

Sequencing of the genome of Ficus carica showed a genome size estimated at 333
Mbp, of which 80% is anchored on 13 chromosomes, very rich in repeated
sequences, including LTR-RE (retro-transposons with long terminal repeats) and
which are the major constituent of the whole genome (Vangelisti et al. 2021).
LTR-REs are mobile DNA sequences, and this characteristic gives them a vital
role in producing an immense diversity of expression and gene function in plant
species and consequently leads to a significant contribution to the evolution of flora
(Lisch, 2013). Indeed, they can promote chromosomal rearrangements by allowing
illicit recombination.
(Devos et al., 2002) through distant locations on the same or other chromosomes,
its coding sequence, or splicing model (Dubin MJ et al., 2018).
In the same context, a study conducted by Usai and his research team in 2019
aims to highlight the abundance of truncated trees in the fig tree genome. In this
study, the entire genome was scanned using predictive tools based on structure and
homology, and they were able to identify a total of 123.8 Mbp of repeated sequences,
representing 37.3% of the genome assembly (Fig. 4.1). TEs were the most abun-
dant, covering 33.57% of the assembly (11.06 Mbp), while tandem repetitions rep-
resented 3.82% (12.74 Mbp). The most abundant TEs are retrotransposons or
PCR-RFLP fingerprinting.
94 R. Ben Abdallah et al.

Fig. 4.1 Annotation of the fig genome. Pie chart showing the percentage of genes (i.e. protein-
coding genes, rRNAs, tRNA and ncRNAs) (left), repeats (i.e. LTR Retrotransposons, Non-LTR
Retrotransposons, DNA transposons and tandem repeats) (right) and unknown sequences. (Usai
et al., 2020)

The rapid and precise identification of various plants was made possible by
molecular fingerprinting employing genes and non-coding areas. Specific primers
can be created by aligning the separated nucleotide sequences, allowing for the
identification of high variation in genes and areas. A PCR product can also be frag-
mented by restriction enzymes, resulting in fragment length polymorphisms. RFLP-
PCR analysis has been regarded as a good tool for distinguishing closely related
genotypes and a quick and accurate method for plant identification.
RAPD (Random Amplified Polymorphism DNA) is a PCR reaction in which the
experimenter does not choose the amplified DNA segments but is amplified “at
random”. This tool has been a prime tool for considering fig genetic variability in
various studies. Khadari et al. (1994), in their research work entitled Varietal
Identification and Genetic Resources in Fig (Ficus carica L.) using RAPD markers,
succeeded in identifying 21 figs (Ficus carica L.) samples representing different
cultivated varieties. In this study, 19 markers were obtained using 12 primers, mak-
ing it possible to characterize 17 genotypes. The primers used revealed a significant
polymorphism. This makes these markers an effective tool for varietal identification
in fig trees.
They could be used to analyze the genetic diversity of more cultivated varieties
and natural populations of Ficus carica. According to another study conducted by
Ikegami and his research team in 2009, which used more than one molecular marker
among other ISSR, RAPD, and SSR. The results obtained suggest that the genetic
diversity of the fig population is not too great; this requires the use of several molec-
ular markers to estimate the fig kinship at the level of the variety. In addition, it was
assumed that ‘Houraihi’, the oldest variety in Japan, was distributed independently
of other foreign varieties in the seventeenth century or earlier.
Indeed, each marker system produced incompletely separated clusters, although
a weak linkage group based on race type appeared in the combined data set. In addi-
tion, Molecular Variance Analysis (AMOVA) showed that most of the total poly-
morphism (Fig. 4.2) was attributable to intra-group variance (ISSR + RAPD, 97.4%;
SSR, 90.1%).
4 Genetic Diversity of Fig Varieties 95

Fig. 4.2 Dendrogram generated using UPGMA; the method used a simple matching similarity
coefficient estimated on 19 fig varieties based on combined ISSR, RAPD, and SSR data (227mark-
ers). Geographic origins are given with the following abbreviations: Japan (J), USA (U), France
(F), and Spain (S). The scale indicates the genetic distance. (Ikegami et al., 2009)

2.4.2 SSR Markers

The microsatellites consist of two-, three- or four-nucleotide sequences repeated in

tandem (always in the same direction). They are uniformly distributed in several cop-
ies over the whole genome of a species and have a high polymorphism rate. This
polymorphism is based on the number of repetition units constituting the microsatel-
lite variation. It is a molecular analysis tool of choice for studying genetic diversity,
especially in plant species, since they are an essential component of their genomes and
are distributed throughout the genome (Deng et al., 2016). Several studies have been
based on SSR as a powerful molecular tool to elucidate genetic variability fig. Among
other things, the work of Boudchicha, in 2019, in the context of her thesis, which was
based on molecular characterization by the use of 22 SSR-type molecular markers on
34 cultivars represented by 77 fig accessions made it possible for the first time to
remove the veil on the state of the diversity of this species in Algeria.
Unlike another study carried out by Essid and her research team in 2015, which
aimed to Analyze the genetic diversity of accessions of Tunisian caprifig (Ficus
caric L.) using simple sequence repeat markers (SSR). However, the results of this
study showed a low genetic diversity. Indeed, even though the sampling sites were
very diverse and distributed over different territories, no clear grouping based on
geographical origin is observed, suggesting a generalized exchange of caprifig plant
material by vegetative multiplication between different areas of the Tunisia territory.
In the same context, another study conducted by Saddoud and his collaborators
in 2005 used microsatellites to characterize 16 cultivars (Ficus carica L.). This
study suggests a high polymorphism rate: 4 to 12 alleles per locus, and mean hetero-
zygosity of 0.656 were noted. The resolution power (Rp) of the six microsatellites
tested ranged from 2.12 to 3.87 for the 16 cultivars studied, showing significant
96 R. Ben Abdallah et al.

genetic diversity (Ht = 0.762). According to this study, the characterization of acces-
sions belonging to different varieties was possible, showing the power and effective-
ness of the molecular tools used.

2.5 Biochemical Traits and Phytochemical Contents Divers

Plant secondary metabolites, often known as phytochemicals, are non-nutritive

plant metabolites that are crucial for plant survival, healthy growth, and reproduc-
tion. Most of these components have biological properties that regulate animal bio-
chemistry and metabolism and have the potential to impact human health. The
leaves of the Ficus carica have the greatest diversity of compounds, with the largest
density of all compounds (except aldehydes and monoterpenes), trailed by the pulps
and peels of the fruits (Barolo et al., 2014; Salem et al., 2013). Numerous bioactive
substances, for example, phenolic compounds, anthocyanin structures, triterpe-
noids, coumarins, phytosterols, organic acids, and volatile composites such as ali-
phatic alcohols, hydrocarbons, and several more chemicals, were identified in
phytochemical research on Ficus carica L. secondary metabolites from several por-
tions of the Ficus carica L. (Fig. 4.3). The majority of fig species include phenolic
composites, organic acids, and volatile complexes. The volatile compounds found
in the fruits (Trad et al., 2012; Mawa et al., 2013; Ware et al., 1993; Gibernau et al.,
1997) and leaves (Mawa et al., 2013; Wang et al., 2017) of F. carica are classified
as terpenes (monoterpenes and sesquiterpenes), alcohols, aldehydes, ketones, esters,
and some other compounds. Terpenes, including monoterpenes (C10) and sesquiter-
penes (C15), are the most common plant secondary metabolites. In typical atmo-
spheric circumstances, these chemicals’ high vapor pressures allow considerable
escape into the air (Trad et al., 2012). Monoterpenes like linalool and, more promi-
nently, epoxylinalool are linked to the fig/wasp linkage (Trad et al., 2012), where
they play a crucial role in attracting certain pollinators. Although sesquiterpenes
made up only 3% of total volatiles in Tunisian cultivars, they were the most com-
mon chemicals found in leaves, with germacrene D, β-caryophyllene, and τ-elemene
being the most common (Barolo et al., 2014). Different portions of the F. carica
were found to contain prevalent terpenes.
α-Pinene, one of the principal monoterpenes reported in many studies, has only
been detected in fruits, while sesquiterpenes have only been found in monovarietal
fig spirits from Portugal (Rodríguez-Solana et al., 2018). Fruit and leaves include
common chemicals such as menthol, τ-muurolene, and τ-cadinene [2, (Barolo et al.,
2014). The more advanced chemical groups in matured fruits include alcohols,
esters, and ketones, which account for 41% of the total aroma.
Different alcohols were identified in fruits [(Z)-3-hexen-1-ol], leaves [2-methyl-
1-butanol and 1-heptanol], spirits [methanol, ethanol, 1-propanol, 1-butanol, isobu-
tyl alcohol, 1-hexanol, octanol, and decanol, among others], in both fruits and leaves
[1-penten-3-ol, benzyl alcohol, and (E)-2-nonen-1-ol] (Mawa et al., 2013; Barolo
et al., 2014), leaves and spirits [1-heptanol] (Barolo et al., 2014; Rodríguez-Solana
4 Genetic Diversity of Fig Varieties 97

Fig. 4.3 Chemical structures of compounds from Ficus carica. (Mawa et al., 2013)

et al., 2018), and lastly in raw materials then spirits [3-methylbutanol, phenylethyl
alcohol] (Mawa et al., 2013; Barolo et al., 2014; Miličević et al., 2017). It is essen-
tial to remind that the production process’s processing conditions influence the con-
centration. Since this component is naturally present in fruits, higher concentrations
were detected in spirits made from fresh figs, while lower amounts were identified
in spirits made utilizing immobilized yeast cell technology (Miličević et al., 2017).
98 R. Ben Abdallah et al.

Heptanal, octanal, nonanal, 2-methyl-butanal, (Z)-2-heptenal, (E,E)-2,4-heptadienal,

(E)-2-octenal, and (E,Z)-2,6-nonadienal are the aldehydes found solely in fruits
(Mawa et al., 2013; Barolo et al., 2014). The ketone 3-hydroxy-2-butanone (acet-
oin) was the first significant chemical discovered in non-pollinated and pollinated
figs (Trad et al., 2012). 6-Methyl-5-hepten-2-one and 3-pentanone were found in
the fruits (pulps and peels) and leaves of Portuguese fig cultivars, respectively
(Mawa et al., 2013; Barolo et al., 2014).
Esters are essential to fruit aroma, especially in ripe figs. They are made by
esterifying alcohols and acyl-CoAs obtained from fatty acid and amino acid metab-
olism, mediated by the enzyme alcohol-o-acyltransferase (Trad et al., 2012). In non-
pollinated fruits, these chemicals are substantially less developed, and their amount
diminishes when immobilized cells are used during the fermentation process.
Esters, notably fatty acid ethyl esters such as ethyl decanoate, ethyl octanoate, and
ethyl dodecanoate, make up the greatest class of volatiles in spirits (95 percent of
whole volatiles). Butyl acetate and isoamyl acetate with a banana odor were the
second and third significant chemicals found in Tunisian kinds of fruits (Trad et al.,
2012). Ethyl salicylate was also another ester found in fruits. Leaves included
methyl butanoate, hexyl acetate, and ethyl benzoate, while fruits and leaves con-
tained methyl hexanoate besides methyl salicylate, identified in fruits, leaves, and
spirits.
The shikimic, malic, oxalic, fumaric, and citric acids were recovered from the
fruits and leaves of F. carica (Mawa et al., 2013; Barolo et al., 2014; Salem et al.,
2013; Soni et al., 2014), but the quinic acid was only found in the leaves (Mawa
et al., 2013; Barolo et al., 2014). Most plant foodstuffs include phytosterols, with
vegetable oils getting the highest quantities. Sterols (modified triterpenes) such
as -sitosterol (Salem et al., 2013), as well as the triterpenoids methyl maslinate,
oleanolic acid, taraxasterol, w-taraxasterol ester, calotropenyl acetate, bauerenol,
24-ethylenecycloartanol, lupeol, and lupeol acetate, have been found in fig leaves
(Mawa et al., 2013; Barolo et al., 2014), betulinic acid in fruits (Wojdyło et al.,
2016), while stigmasterol was reported in both (Soni et al., 2014; Joseph & Raj,
2011). Polyunsaturated fatty acids made about 84 and 69% of total fatty acids in
dried and fresh F. carica fruits. Linoleic acid detected in fresh and dried fruits was
the only polyunsaturated fatty acid detected in fresh and dried fruits. Considering
monounsaturated fatty acids, the most prevalent in fruits is oleic acid (Barolo et al.,
2014). The fig fruit and bark of F. carica provided fifteen anthocyanin pigments.
Most of them have cyanidin as an aglycone and pelargonidin derivatives (Ahmad
et al., 2013). Overall, individual phenolic compounds, including phenolic acid,
chlorogenic acid, flavones, and flavonols, were extracted from fresh and dried fig
skins of F. carica.
Dried figs had total greater levels of phenolics than fresh fruit pulp, owing to the
input of the dry skin. The main significant phenolic was quercetin rutinoside (Soni
et al., 2014), while microbial β-D-glucans were obtained from Libyan figs of F. car-
ica (Joseph & Raj, 2011). The pulps and peels of figs were used to extract phenolic
acids such as 3-O- and 5-O-caffeoylquinic acids, ferulic acid, quercetin-3-O-gluco-
side, quercetin-3-O-rutinoside, psoralen, and bergapten, as well as organic acids
4 Genetic Diversity of Fig Varieties 99

such as oxalic, citric, malic, shikimic, and fumaric acids (Salem et al., 2013). Many
biological activities have been considered and validated on F. carica extracts, and
bioassay-guided fractionation has, in most cases, permitted the chemical structures
involved for such biological effects to be assigned, finalizing much of its traditional
applications (Barolo et al., 2014). Phenolic compounds are among the most signifi-
cant phytochemicals investigated in F. carica, having an antioxidant activity (AC).
Many of these chemicals can operate as antioxidants in various ways, including
reducing agents, hydrogen donors, free radical scavengers, and singlet oxygen
quenchers (Mawa et al., 2013). In ethanol (40%) leaf extracts of F. carica (solid to
liquid ratio 1:60 g/mL, temperature extraction of 60 °C, and 50 minutes of ultra-
sonic treatment), the highest total flavonoid content (25.0 mg/g) with noticeable
scavenging activities against hydroxyl and superoxide anion free radicals in a
dependent way were reported (Ahmad et al., 2013).
Different studies have looked at the AC of fruit extracts. Total polyphenolic con-
tent (TPC) and total flavonoid content (TFC), as well as the quantity and pattern of
anthocyanins, were measured in extracts from six commercial fig types using the
ferric reducing antioxidant technique (FRAP). TPC, TFC, and anthocyanins (cyan-
idin-3-O-rutinoside as the major constituent) were found in abundance in the sam-
ples with the greatest AC (Mawa et al., 2013; Ahmad et al., 2013). Using in vitro
scavenging properties on DPPH⋅, superoxide, and hydroxyl radicals as well as
reducing power tests, two fruit extracts [water (WE) and crude hot water-soluble
polysaccharide (PS)] were assessed for AC in another study. Both extracts demon-
strate significant DPPH⋅ scavenging activity [WE (EC50, 0.72 mg/ml) and PS (EC50,
0.61 mg/ml)], while PS has the greatest superoxide radical scavenging activity
(EC50, 0.95 mg/ml) and the hydroxyl anion radical (43.4% at 4 mg/ml concentra-
tion) (Ahmad et al., 2013). Holm oak acorns, in addition to carob pods, were com-
pared to ethanolic extracts from white Beni Maouche Algerian figs
(Amessis-Ouchemoukh et al., 2017).
Fig extracts had lower DPPH⋅ scavenging efficacy (20.5%) than ABTS radicals
(68.9%) but better phosphomolybdenum lowering ability (638 mg Gallic Acid
Equivalent, GAE, per 100 g). This extract (73.1%) also prevented the generation of
the Fe2 + −ferrozine complex and effectively scavenged H2O2. The nitric oxide
(NO) radical scavenging capacities of the extracts from the three studied (carob,
acorns, and figs) fruits were not significant (Amessis-Ouchemoukh et al., 2017).
Chemiluminescence utilizing lucigenin was used to assess the generation of reac-
tive oxygen species (ROS) in the presence of ethanolic fig extract. After 15 minutes
of treatment, the extract at the highest concentration (250 μg/mL) appeared to
achieve its higher level of lucigenin inhibition, with a value 44% lower than that
procured with diphenylene iodonium (0.2 mM), the standard preferential inhibitor
of nicotinamide adenine dinucleotide phosphate (NADPH) oxidase tested (Amessis-
Ouchemoukh et al., 2017).
The enzyme xanthine oxidase (XO), which produces reactive oxygen species,
was inhibited by ethanolic fig extracts. As a result of its prooxidant impact, the
extracts evaluated at 500 μg/mL exhibited a reduction in the inhibition of XO activ-
ity. The high correlation coefficients between XO inhibitory activity and phenolic
100 R. Ben Abdallah et al.

substances and flavonoids show XO’s inhibition activity. The equilibrium between
the synthesis of reactive oxygen species (free radicals) and antioxidant defenses are
disrupted in oxidative stress, resulting in the formation of reactive oxygen species
(ROS). Four groups: Streptozotocin-induced diabetic rats, diabetic rats given a sin-
gle dose of a basic fraction of F. carica leaf extract, diabetic rats given a single dose
of a chloroform fraction of the extract, and normal rats were used to study the role
of these free radicals in the creation of tissue destruction in diabetes mellitus.
Diabetic animals had increased erythrocyte catalase activity, vitamin E plasma lev-
els, monounsaturated and polyunsaturated fatty acids, saturated fatty acids, and lin-
oleic acid levels than control animals. All F. carica fractions were shown to correct
the fatty acid and plasma vitamin E levels of diabetic animals. The isolation of a few
groups of plant metabolites has resulted from phytochemical studies on F. carica.
Most phytochemical studies on F. carica have focused on the leaves and fruits, with
little knowledge of the stem and root phenolic profiles. However, given the wide
range of traditional applications and well-established pharmacological effects of
F. carica, there is still unlimited attention for phytochemical research employing
bioassay-guided isolation. Future studies in the areas above will provide convincing
support for the future therapeutic usage of F. carica in modern medicine.

2.6 Proteomic and Transcriptomic Analysis

Multi-omics provides data integration and processing to acquire insights into the
interrelationships, functioning, and biological processes at several levels of biologi-
cal systems (Fabres et al., 2017; Padden et al., 2014). In recent years, multiple
research projects on the transcriptome elements of fig fruit growth and ripening
have been conducted. Transcriptomic comparison of young San Pedro type fig and
common fig fruit revealed that zeatin biosynthesis and plant hormone signaling
pathways are differentially regulated (Chai et al., 2017), and ethylene synthesis and
phytohormone signaling were found to be differentially expressed between caprifig
and common fig fruit (Ikegami et al., 2013). Fruit cell wall-modification enzymes,
genes encoding ethylene-response factors (ERFs), and ascorbate oxidase were sig-
nificantly increased during fig ripening at the transcriptome level (Freiman et al.,
2014). The final biofunction executor is the protein, the end outcome of gene expres-
sion. Proteomic research of essentially biological processes, including fruit growth
and ripening, may give more detailed insights (Palma et al., 2011). Isobaric tags for
relative and absolute quantitation (iTRAQ) are a commonly utilized quantitative
method in proteomics investigations that enhances protein quantification accuracy
and reliability over traditional 2D-polyacrylamide gel electrophoresis (PAGE)
(Miyagi & Rao, 2007). As a result, it might help reveal and speculate on the exten-
sive and widespread alterations that underpin critical fruit ripening, quality determi-
nation, and horticultural trait creation, making commercial and market-driven fig
quality management easier for innovation.
4 Genetic Diversity of Fig Varieties 101

In recent work, researchers used BLASTN (Basic Local Alignment Search Tool
for nucleotide sequence analysis) algorithm to compare the projected transcrip-
tomes of two fig cultivars: the Italian cv Dottato and the Japanese cv. Horaishi
(Raskovic et al., 2016). While most genes were discovered in both cultivars, a small
number of genes were detected only in the Japanese or Italian cultivars.
Phosphoglycerolipid metabolism, which is involved in membrane composition and
signal transmission, and cyano amino acid metabolism, which is engaged in chemi-
cal defense against herbivores and pathogens, were considerably over-represented
in the cv. Dottato projected transcriptome Among Kyoto Encyclopedia of Genes
and Genomes (KEGG) Pathways. For RNA-seq gene expression investigations, the
availability of a gDNA-based reference transcriptome is the ideal choice. For exam-
ple, in tree species under abiotic stress, such a transcriptome was applied (Fabres
et al., 2017; Sun & Hu, 2016). Such reference transcriptome is available for fig
(Raskovic et al., 2016). The availability of a genotype‘s expected transcriptome
allows for a more accurate and thorough examination of gene expression in that
genotype. Furthermore, increasing the number of reference transcriptomes for a
species allows for the characterization of that species’ pan-genome. The fig refer-
ence transcriptome had 41,857 predicted genes from F. carica cv. Dottato.
Gene ontology and metabolic pathways were used to classify predicted genes.
There were disparities in the cultivars‘anticipated gene repertoires, with 4803 and
2383 genes discovered in the Horaishi and Dottato predicted transcriptomes. Several
genes are unique to the cv. Dottato projected transcriptomes are involved in chemi-
cal resistance against herbivores and pathogens, which is essential. Those findings
provide a resource for fig functional genomics and may be used to answer issues
about growth, protection, and environmental adaptability in this plant species.

3 Multivariate Statistical Analysis of Qualitative

and Quantitative Traits

The analysis of the joint behavior of more than one random variable is done using
multivariate statistical approaches. Multivariate approaches come in a variety of
forms. Multivariate statistical methods have been employed in several research on
the genetic diversity of figs. A study was conducted to assess the genetic variability
of a fig tree (Ficus carica L.) gene bank in the Tunisian Sahel region (Chatti et al.,
2004). It was based on morphological characteristics related to the tree’s vegetative
development. The data obtained were subjected to various statistical analyses using
SAS (Statistical Analysis System) software: principal component analysis (PCA)
and canonical discriminant analysis, providing the Mahalanobis phenotypic dis-
tances for the parameters taken together. A classification dendrogram was estab-
lished from these distances to translate the phylogenic relationships between the
cultivars studied. Means were compared using Duncan’s test.
102 R. Ben Abdallah et al.

Pereira et al. (2015) investigated the agronomic behavior and quality of six fig
cultivars for fresh consumption grown in Extremadura, Spain (the breba and major
crops). SPSS for Windows, version 19.0, was used to do a statistical analysis of the
data. Multivariate analysis of variance was used to investigate the mean values of
the interaction factors ‘cultivar-year,’ ‘cultivar-block,’ and ‘year-block’ for pomo-
logical and quality aspects (ANOVA). In addition, Tukey’s honestly significant dif-
ference (HSD) test (p ≤ 0.05) was used to examine mean values.
The principal phenolic components, phenolic profiles and antioxidant activity in
nine sun-dried fig cultivars with varying skin colors originating from South-Eastern
and Middle-Eastern Tunisia are reported (Khadhraoui et al., 2019). Statistical anal-
yses were carried out using an ANOVA with SPSS version 22.0 software. The
Duncan test was used to assess significant differences, with a significance level of
p < 0.05, and Pearson’s correlation coefficient (r) was employed to determine cor-
relations between analyzed parameters. Principal component analysis was used to
examine the structure of genetic variability across fig cultivars based on mean val-
ues of total polyphenol content, phenolic compounds such as phenolic acids and
flavonoids, fatty acids, and antioxidant activity (PCA). To examine the role of phe-
nolic chemicals in the differentiation of Tunisian dried fig samples, these multivari-
ate analyses were carried out using the Xlstat 9.0 software.
The morphological and pomological variability of edible fig genotypes belong-
ing to the Smyrna-type was investigated (Khadivi et al., 2018) from the Estahban
region in Iran’s Fars province. For statistical analysis, the average values of pheno-
typic data were employed. The mean, minimum, maximum, standard deviation, and
coefficient of variation for the measured features were calculated. SAS software
was used to assess the variance for all of the characters. SPSS software was used to
calculate Pearson correlation coefficients between the attributes. Using the SPSS
software, the correlations between the genotypes were determined using principal
component analysis (PCA). Using PAST statistics software, a distance matrix cre-
ated from phenotypic data was used for Ward method cluster analysis to understand
the patterns of variability among genotypes better.
Ahmed et al. (2015) characterized 71 farmed, and wild Tunisian fig trees using
eight morphological features and 17 simple sequence repeats loci. Statistical evalu-
ation of physical characteristics, the principal component analysis (PCA) was used.
A phenetic analysis was conducted using the coordinates of the three first axes of
this PCA. The function “hclust” was used to do hierarchical clustering using the
unweighted pair-group approach of arithmetic averages (UPGMA). The Kelly
Gardnere Sutcliffe penalty function for a hierarchical cluster tree was calculated
using the “kgs” package maptree, Version 1.4–7, and a dataset with several clusters.
The distance (or similarity) level at which two observations become part of the same
cluster is defined as the cophenetic distance between them.
4 Genetic Diversity of Fig Varieties 103

4 Variation in Graft Compatibility of Wild Fig Species

as Rootstocks

Grafting is a technique that has been mastered for a long time to cultivate fruit trees
and conifers. It was known at least six thousand years BC in the Far East. When we
study its long history, we discover that grafting was applied even to herbaceous
plants both in China and ancient Greece, far from being limited to fruit trees.
Nevertheless, grafting is an essentially conservative operation. This opinion was
established little by little because most of our fruit varieties are hybrids, which
result from a crossing of one or more other forms and whose qualities cannot be
reproduced by sowing their seeds. Thus, many improved varieties have also become
incapable of reproducing other than by vegetative means because they no longer
even give fertile seeds (Pierre, 1959).
In other contexts, grafting has become an indisputable solution to solve problems
for certain fruit varieties which are a severe threat to their crops because of several
factors such as soil disturbances, for example, the presence of nematodes (Condit,
1933), soil disease (Hosomi, 2015) and ceratocyst cancer (Figueiredo & Pinheiro,
1969); therefore rootstocks resistant to such soil disorders have long been desired.
Some F. carica rootstock cultivars are adapted to grow under these unfavorable
conditions (Valarini & Tokeshi, 1980; Hosomi, 2015). Nevertheless, these root-
stocks are not immune and cannot completely prevent the adverse effects of such
soil disorders. In this context, the work of Hosomi and his collaborators in 2015
takes on its value in trying to assess the compatibility of grafts. Their study con-
cluded that A possible compatibility of the common fig tree scion was anticipated
on the rootstocks of F. palmata, F. septica, F. pumila, and F. stipulata. These root-
stocks have previously been classified in the same subgenus as Ficus (Corner, 1965),
and the observed compatibility probably reflects a taxonomic relationship.
Interspecific hybridization of common fig trees and wild Ficus species has been
attempted to create a truly resistant rootstock (Yakushiji et al., 2012). According to
their results, evaluating the compatibility of species grafts from the various taxo-
nomic groups would help identify the best possible cross parents for hybridization.
In the same context, another study was carried out by Yakushiji and his collabora-
tors in 2012 to seek a solution to Ceratocystis canker disease. It is a severe disease
of the fig tree (Ficus carica L.) which can even kill the tree, caused mainly by the
soil fungus Ceratocystis ficicola. During this study, crosses were made, and they
concluded that the backcrossing of F. carica with the interspecific hybrid between
F. carica and F. erecta contributes to establishing practical fig tree rootstocks with
resistance Ceratocystis canker.
104 R. Ben Abdallah et al.

5 Trends in Fig Germplasm Conservation

For centuries, fig germplasm has been damaged by severe genetic loss. As a result,
it is essential to build up programs worldwide to restore and typify fig genetic
resources, and such initiatives have evolved in many countries. As a result, fig germ-
plasm enlistments have been documented in Asia, Europe, North Africa, and North
America. With 442, 314, 190, 180, and 154 accessions, respectively, Italy, Turkey
(Elbreyli Fig Research Institute), California (National Clonal Germplasm Repository
(NCGR) Davis), Turkmenistan (TES-PGR), and France (CBNM Porquerolles) have
the greatest collections. In addition, distinct fig cultivars (154) have been gathered
from Italy, North Africa, Greece, Turkey, France, Tunisia, and Spain for the CBNM
Porquerolles collection.
There are 190 accessions in the NCGR fig gathering. The Institute of Arid
Regions of Medenine, Tunisia’s Arid Land and Oasis Cropping Laboratory germ-
plasm collection contains 119 native fig accessions and 20 caprifigs gathered from
Tunisia, Algeria, and Egypt. In situ or on-farm conservation of fig cultivars in
Tunisia is an issue that research institutes, local nongovernmental organizations
(NGOs), development agencies, and farmers are tackling. Specific geographic sites
were chosen on the way to protect and transplant native fig varieties. In southern
Tunisia, the Zammour and Douiret districts were designated traditional places to
conserve fig varieties (Mars et al., 2008). The evolving climatic circumstances
linked with climate change have become deeply critical in vitro conservation of fig
germplasm. Tissue culture is used to maintain genomic resource collections in vitro
(IPGRI & CIHEAM, 2003).
Cryopreservation and the slow-growth approach are two in vitro conservation
procedures that have been established (Engelmann, 2000). Cryopreservation is the
preservation of germplasm in liquid nitrogen at 196 °C for long-term storage; cel-
lular divisions and metabolic functions are halted at such temperatures. Genetic
instability, on the other hand, poses a challenge to preserved plant materials (Panis
et al., 2000). For cryopreservation of fig materials, two procedures are used: the
traditional technique and the vitrification technique (Engelmann, 2000). The tradi-
tional approach entails two steps: gradual controlled chilling followed by rapid
immersion of plant components in liquid nitrogen. A penetrating cryoprotectant,
such as dimethyl sulfoxide, ethylene glycol, or polyethylene glycol, is used to treat
samples in this approach. This method keeps water from crystallizing inside cells
and preventing cell damaging (Panis et al., 2000).
Furthermore, traditional cryopreservation is inefficient for fig tissue such as cal-
lus and cell suspension cultures. The vitrification approach was used to preserve
plant tissue cultures and is the most commonly used in the plant cryopreservation
procedures (Panis et al., 2000). Plant components such as shoot tips, buds, and meri-
stem are cultivated under freezing temperatures in the slow growth technique, which
keeps the plant material alive but drastically slows the growth (Engelmann, 2000).
The slow-growth approach is the most effective for preserving shoot tips in vitro.
Treatment of shoot tips with cryoprotectant for 20 minutes and subsequently
4 Genetic Diversity of Fig Varieties 105

chilling to 40 °C resulted in a 50% survival rate for plant regeneration. The most
frequent method for cryopreservation of shoot tips, meristems, cell cultures, and
somatic embryos is slow growth. Plant regeneration from cryopreserved root tips
was produced by adventitious bud development; however, no difference was identi-
fied between the cryopreservation and slow-growth procedures for root tip explants
(Towill & Bajaj, 2002).

6 Conclusion

Understanding the genetic basis of the complete metabolic system involved in cap-
rification will be critical for developing cultivars with greater yield and longer shelf-
life products. Findings show that when studying genetic diversity in fig trees, two
methodologies should be considered: morphological and molecular. Molecular
analyses should be used in conjunction with morphological and horticultural evalu-
ations because they give additional information, improve the resolving power of
genetic diversity investigations, and help understand the domestication process.
This has crucial consequences for fig management, necessitating strategies to ensure
the species’ durability and genetic variety. In addition, knowing about the genetic
diversity in this germplasm will make it easier to employ in breeding operations and
enhance its management in the face of climate change.

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Chapter 5
Bud Structure and Evolution

Giuseppe Ferrara and Andrea Mazzeo

1 Fig Buds Description

In temperate climates, the buds of deciduous fruit trees are dormant during the
autumn and winter when the coldest temperatures occur. It is commonly assumed
that this rest period consists of an endo-dormancy phase, followed by an eco-
dormancy phase, and finishes with para-dormancy during the growing season (Lang
et al., 1987; Crabbe, 1994; Faust et al., 1995, 1997). Para-dormancy is synonymous
with correlative inhibition as manifested in apical dominance (AD) (Crabbe &
Barnola, 1996) and occurs during the vegetative/reproductive phases in the growing
season. In contrast, endo-dormancy requires a cold period to release buds from an
‘inactive’ stage. Apart from overcoming AD, lateral buds still have to meet their
specific chilling requirement (Faust et al., 1995). Terminal buds are generally easier
to release from dormancy than auxiliary buds (Crabbe, 1994). The fig tree (Ficus
carica L.) is one of the oldest perennial fruit species cultivated in the Mediterranean.
According to Eisen (1901), Ficus carica (common fig) is a deciduous species in
most of the world (Fig. 5.1) but is evergreen in tropical areas (Condit, 1947). The
common fig produces both vegetative and flower buds at the axil of the leaves
(Fig. 5.2) in summer-fall, and some of them evolve into fruits during the current
season (main crop), and others could overwinter (breba crop in the successive sea-
son or dormant and latent mixed/vegetative buds). Vegetative buds are generally
smaller, thinner, and conical (Fig. 5.3), whereas the flower buds are larger and
rounded (Fig. 5.4), although often the difference is relatively small (Condit, 1947;
Sarkhosh et al., 2022). The terminal bud is a mixed bud, which is larger and longer
than the other two, conical, and has an attenuate tip (Fig. 5.5). This bud develops in

G. Ferrara (*) · A. Mazzeo

Department of Soil, Plant and Food Sciences, University of Bari “Aldo Moro”, Bari, Italy
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 109
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_5
110 G. Ferrara and A. Mazzeo

Fig. 5.1 Fig orchard in July (a) and during winter rest (b)

spring, producing leaves and flower buds, either producing fruit (main crop), or
overwintering flower and vegetative buds. Sometimes sub-apical or lateral buds are
also mixed buds.
Unlike many temperate species that have a single blooming season and differen-
tiate the buds, for the successive spring flowering, during the previous summer or
autumn, the common fig continually produces flower buds during the growing sea-
son (Fig. 5.6). Fig buds require little or no winter chilling to break endo-dormancy
(Ferguson et al., 1990), and growth resumes in early spring (Flaishman et al., 2008).
Under temperate growth conditions, with no limiting water and nutrient conditions,
the common fig produces several flower buds during the growing season until the
autumn developing into fruits (main crop), or these buds then overwinter and pro-
duce a first spring fruit crop (called brebas or breba figs). The brebas crop may be
destroyed in cold climates by late spring frosts (Fig. 5.7). The brebas are borne on
the previous year or “old” wood, the 1-year shoot, whereas the second fruit crop or
main crop is borne on the shoot grown during the current season (Fig. 5.8). The
leaves fall at the end of the growth period, and the tree enters the endo-dormancy
5 Bud Structure and Evolution 111

Fig. 5.2 Developing fruits (from flower buds) and vegetative buds at the axil of the leaves of
the shoot

Fig. 5.3 Small and thin vegetative buds between two developing fruits

period. The length of the dormant season and the budburst time in spring depend
upon local climatic conditions. Due to strong apical dominance in many cultivars,
the seasonal growth is either from the terminal bud or the adjacent one or two buds
(Fig. 5.9), even though apical dominancy varies depending on the cultivar (Ersoy
et al., 2008).
The number and sizes of flower buds are closely related either to the vegetative
growth of the tree or to the size of the crop maturing in summer and fall. Caprifig
trees, which have a light summer and fall crop, commonly produce an enormous
number of flower buds that grow in the spring and mature a crop in June, the profichi
112 G. Ferrara and A. Mazzeo

Fig. 5.4 Longitudinal section of a rounded fruit bud. Measures are indicated in μm

Fig. 5.5 Longitudinal

section of a conical
terminal mixed bud

crop (Condit, 1947). The color of the fruit bud is approximately similar to that of the
terminal mixed bud, from greenish to reddish. The color of the terminal bud is
largely correlated with the color of the fruit, most green-fruited cultivars have green
buds, and most dark-fruited cultivars have buds with the shade of red-violet color
(Fig. 5.10). Some cultivars having dark figs, such as Col de Dame Noir and Gouraud
Rouge, show green terminal buds; vice versa, some with green figs, notably Genoa,
have colored buds. All the fig buds are developed on the same shoot, i.e., fruit
(flower) bud, vegetative bud, and mixed bud. In the common fig, the basal portion
of the shoot holds flower buds evolving into the main crop, whereas the distal part
of the shoots holds the overwintering flower buds developing (or not) the brebas in
the successive spring. The buds can be isolated but often are aggregated with two
5 Bud Structure and Evolution 113

Fig. 5.6 Main crop fruits developing along the growing shoot

Fig. 5.7 Buds injured by

late spring frost. Necrosis
is visible on flower,
vegetative and mixed buds

flower buds and a vegetative bud in the middle as in the cultivar Sultani (Fig. 5.11)
or only a flower bud and a vegetative bud or even one/two flower bud (Fig. 5.12).
When two flower buds and one vegetative bud are found in the axil of each leaf,
rarely more than one of the flower buds ever develops into a fig in the Mission
114 G. Ferrara and A. Mazzeo

Fig. 5.8 Presence of both

brebas (bigger and on the
1-year shoot) and main
crop fruits (smaller and on
the current year shoot)

cultivar, and the remaining bud generally remains latent, while in Kodota and
Calimyrna or some other cultivars it is quite common for two figs in the axil of an
individual leaf to develop to maturity (Petrucci & Crane, 1950). Some fig cultivars
also have flower buds in the distal portion, which may develop in late summer-
autumn into the fruit as a third crop (or even the only main crop), in some areas
defined as the late main crop (Christmas or winter fruits).
The fig bears numerous latent buds, generally protected by tick scales, abundant
on 2–3-year-old branches, and which sometimes can burst and produce fruits but of
poor quality (Grassi, 1991); these latent buds are often permanently arrested in their
development and never burst.
The breba or first crop of figs is borne laterally on the wood of the previous sea-
son’s growth from buds produced in the axils of leaves along the distal portion of the
shoot. These buds enlarge the following spring and mature into fruit during the
month of May–June. Morphological and developmental studies conducted by Duric
et al. (1992) during dormancy and growth on cv. Tenica bearing a single crop and
cv. Petrovaca bearing two crops per year reveal that three types of buds occur on
1-year-old shoots. (1) Flower buds which are spherical, have 3–5 scales, with the
5 Bud Structure and Evolution 115

Fig. 5.9 Growth of apical and lateral mixed and vegetative buds. The growing brebas are
also visible

inflorescence axis in the internal cavity enclosing female primordia differentiated to

varying degrees depending on the position of the bud on the shoot. These buds pro-
duce parthenocarpic fruits only in cv. Petrovaca. The number and the size of flower
buds are closely correlated with the vegetative growth and determine the size of the
summer and the fall crops (Flaishman et al., 2008). (2) Mixed buds are conical,
made up of 5–8 scales, and inflorescences on shoots arising from these buds pro-
duce fruit in both cultivars. (3) Vegetative buds, made up of 3–5 scales, usually
remain dormant and may produce shoots in the successive season. The scales cover-
ing the vegetative bud absciss as the bud elongates and develops into a shoot. The
distal buds that do not produce fruit during the current season are in para-dormancy
and overwinter, giving rise to the brebas the following summer.

2 Development and Evolution of Inflorescence

The initiation and development of flower buds was macroscopically examined by

Tanrisever (1998). Primordia, which will develop into inflorescence, were generally
covered with three but sometimes with four scales. Petrucci & Crane (1950) reports
that the first visible evidence of inflorescence differentiation is the broadening and
elongation of the axis and the initiation of scales that eventually surround the osti-
ole of the syconium. The initial 3–4 scales that cover the primordium are persistent
and, upon maturity of the fig, are located at the apex of the stalk. As observed at the
microscope, at the beginning of the growing season, in the cultivar Petrelli, the
116 G. Ferrara and A. Mazzeo

Fig. 5.10 Greenish scales of flower and mixed buds of a cultivar producing fruits with a green skin
(a) and reddish scales of flower and mixed buds of a cultivar producing fruits with a red-violet
skin (b)

Fig. 5.11 Presence of

three buds, two growing
flower buds (main crop)
and a vegetative bud in the
middle (cultivar Sultani)
5 Bud Structure and Evolution 117

Fig. 5.12 Presence of two buds at each node, a single flower bud and a small vegetative bud

Fig. 5.13 Microscopic

observation of a flower bud
in the distal portion of the
current season shoot with
small, short and curved
ostiolar scales and very
small flower primordia
(cultivar Petrelli)

flower buds in the distal portion of the current season shoot have small, elongated,
and curved ostiolar scales with very small female flower primordia (Fig. 5.13).
In contrast, in the basal flower buds, the ostiolar scales were much larger, with
female flowers more developed in the cavity of the syconium (Fig. 5.14) (Marcotuli
et al., 2020). In the Dottato cultivar, differences in the development of the flower
buds were also evident; in the smaller flower buds at the apical position, the ostiolar
scales were still underdeveloped, and the female flower primordia were small and
almost rounded (Fig. 5.15); whereas at the basal nodes the flower buds had ostiolar
scales completely developed and the structures of the female flower primordia were
more evident (Fig. 5.16) (Marcotuli et al., 2020). Further development of the inflo-
rescence primordium consists of a continued broadening of the apex, the cells around
the periphery divide and give rise to many ostiolar scales, and the single flowers
continue to grow, assuming a more cylindrical shape (Fig. 5.17). At this stage of
development, the apex of the axis changes from convex to a concave shape. Following
118 G. Ferrara and A. Mazzeo

Fig. 5.14 Microscopic

observation of a flower bud
in the basal portion of the
current season shoot with
large, elongated and curved
ostiolar scales and
developed female flower
primordia (cultivar Petrelli)

Fig. 5.15 Microscopic observation of a flower bud in the distal portion of the current season shoot
with small, underdeveloped ostiolar scales and very small female flower primordia (cultivar Dottato)

this stage, inflorescence primordia broaden and elongate, forming the neck and stalk
of the syconium. The formation of the syconium was completed when the apical
portion of the cup-shaped structure grows towards the center and forms an ostiole
closed with numerous scales. Buds destined to become flower or vegetative buds are
identical in their initial stages of development. There are generally no visible inflo-
rescence primordia in the mixed buds before bud burst (Fig. 5.18), as observed in
summer for the flower buds, but only an undifferentiated structure until the end of
winter with few exceptions (Fig. 5.19). The mixed bud will first differentiate inflo-
rescence primordia after bursting, and the flower primordia will completely differen-
tiate into the developing syconium for the main crop in spring-summer. In the
5 Bud Structure and Evolution 119

Fig. 5.16 Stereomicroscopic

observation of a flower
bud in the basal portion of
the current season shoot with
large, elongated and curved
ostiolar scales and developed
female flower primordia with
evident ovary and style
(cultivar Dottato)

Fig. 5.17 Microscopic

observation of developing
female flowers in the
receptacle of main crop

distal flower buds, flower primordia will develop into the buds but will rest their dif-
ferentiation during the winter and complete the development in the successive spring
with the production of brebas (Marcotuli et al., 2020).
Flower buds (of brebas) had already formed florets (not completely differenti-
ated) at the beginning of summer, with a receptacle characterized by masses of cylin-
drical pistillate primordia (Fig. 5.20). They remained at this stage from summer until
the end of winter before bud burst (Marcotuli et al., 2020). The X-ray analyses of
flower buds either developing into brebas or the main crop showed these buds very
similar at the first stages of development with the difference only in the number and
thickness of bud scales; the flower primordia in the receptacle are almost identical.
This light difference may suggest that the flower buds of the brebas are prone to
overwinter (number and thickness of scales), whereas the flower buds of the main
crop have lighter scales because they have to develop in the current season and do not
need to overwinter (Marcotuli et al., 2020). In Dottato, the flower buds of the brebas
(Fig. 5.21) and main crop (Fig. 5.22) are almost identical, taken at the same stage.
120 G. Ferrara and A. Mazzeo

Fig. 5.18 Microscopic

longitudinal section of a
mixed bud covered with
scales and the apex is
visible at the bottom
during winter rest

Fig. 5.19 Microscopic

longitudinal section of a
mixed bud covered with
scales and the apex is
visible at the bottom before
bursting. The meristematic
apex and a possible main
crop are also indicated

The flower bud of breba is covered with several bud scales since it is an overwinter-
ing bud. In the case of Petrelli, the two types of flower buds also differentiate for the
more elongated shape and less bud scales of the main crop (Fig. 5.23) with respect to
the brebas (Fig. 5.24) (Marcotuli et al., 2020).
Bud’s position on a shoot about fruitfulness has been investigated, and Petrucci
& Crane (1950) reported that although inflorescence primordia are located at each
node on the shoot, the buds at the distal position of the shoot are less fruitful. The
fruitfulness increases from the third node, where together with nodes 4 and 5, the
main crop is mainly produced. However, fruitfulness strongly depends on other fac-
tors such as crop load, vegetative activity, cultural practices, cultivar, etc. (Ferrara et
al., 2016, 2017; Marcotuli et al., 2019).
The flower bud differentiation begins at the basal nodes at the beginning of the
growing season and proceeds in the successive nodes depending on the cultivar,
climatic conditions, cultural practices, and resource availability (water, nutrients).
The first nodes will differentiate into flower buds and develop the syconium
5 Bud Structure and Evolution 121

Fig. 5.20 Microscopic longitudinal section of a flower bud (brebas) in summer with evident
female flowers primordia

Fig. 5.21 X-ray picture of a small Dottato first crop (brebas) at the beginning of development in
spring. Bud scales, florets and ostiolar scales are clearly evident. (From Marcotuli et al., 2020)
122 G. Ferrara and A. Mazzeo

Fig. 5.22 X-ray picture of a small Dottato main crop at the beginning of development. Florets and
ostiolar scales are clearly evident but the bud scales are much less developed than in the first crop
(brebas). (From Marcotuli et al., 2020)

containing the inflorescences until the ripening of the fruit (main crop). In the distal
nodes, the flower bud differentiation will start during the season but stop at the sum-
mer’s end (Marcotuli et al., 2020). The flower buds of brebas are mainly located in
the distal nodes, a portion of the shoot which grows at the end of the growing sea-
son, as indicated by the shorter length of the internodes, and these buds may be
more susceptible to erratic climatic conditions at the end of summer (rain, cold, heat
waves, etc.). However, as previously reported, flower buds are already differentiated
even at the distal nodes in summer (July–August). In a 3-year study carried out in
Tunisia, the main crop amount exerted a strong influence on the fruiting intensity of
the successive year; in particular, a heavy main crop load decreased the number of
brebas in the following year, while a heavy breba crop load reduced the main crop
load of the current season (Gaaliche et al., 2011) in a sort of productive balance by
the tree for the use of its resources. In the case of a heavy main crop, the number of
flower buds (brebas) overwintering is clearly lower since almost all buds developed
into the main crop during the season. Conversely, the higher number of flower buds
(brebas) retained on one-year shoots and developing in the successive season will
compete with both the current year’s shoot growth and the main crop fruits, thus
reducing both the yield of the main crop and the shoot length (Marcotuli et al., 2020).
The most relevant fruit tree species, such as sweet cherry, apple, peach but also
grape, have flower (mixed) buds which start to differentiate during the summer of
5 Bud Structure and Evolution 123

Fig. 5.23 X-ray picture of a small and elongated Petrelli main crop at the beginning of develop-
ment. Florets and ostiolar scales are clearly visible. (From Marcotuli et al., 2020)

the year 1 (such as the flower buds of brebas), and the process is completed at bud
burst in the successive season for a two-year process (Koutinas et al., 2010). The
distal flower buds (brebas) differentiate in two seasons, whereas the inflorescence of
the main crop differentiates in the same year of formation since, at the end of winter,
only rudimentary primordia are visible in the basal nodes of the apical/lateral mixed
buds (Marcotuli et al., 2020).
Molecular analyses also indicated a higher expression level of APETALA1 and
APETALA2, AGAMOUS genes involved in flower organ formation, in brebas of
both Dottato and Petrelli concerning the main crop. The higher expression of
AGAMOUS (required for stamen development) in breba may suggest a possible
presence of staminate flowers at the ostiole of brebas during its evolution, as the
profichi fruits in the caprifig.
Phylogenetic studies have shown that the common fig has a monoecious ancestor
(Machado et al., 2001) and successively evolved in a gynodioecious species.
Therefore, the presence of these dormant flower buds evolving in brebas at the
124 G. Ferrara and A. Mazzeo

Fig. 5.24 X-ray picture of a small Petrelli first crop (brebas) at the beginning of development in
spring. Florets and ostiolar scales are clearly evident and also the bud scales are well developed.
(From Marcotuli et al., 2020)

beginning of the season may be a relic of the ancient monoecy, with the wasps enter-
ing the different inflorescences (main summer crop, main late crop, breba) during
the year to lay the eggs, as they nowadays do in the caprifigs (mammoni, mamme,
profichi).
Dioecy was considered as an adaptation to seasonal climates in order to support
the Ficus-Blastophaga symbiosis (Kjellberg et al., 1987) or also to stimulate the
inter-crosses to increase the variability of the offsprings.
A study of the genes involved in the natural process of flowering and flower dif-
ferentiation in flower buds (breba or main crop) showed twelve genes expressed in
the two crops (breba and main crop) and in the two cultivars analyzed (Dottato and
Petrelli) involved in the formation of organs of the fig inflorescence (sepals, petals,
stamens, ovary) (Marcotuli et al., 2020).

3 Influence of Cultural Practices and Plant Growth

Regulators (PGRs) on the Buds

The common fig is a species with a strong apical dominance; thus, pruning and
other cultural practices are required to control the growth of leader shoots and pro-
mote the emergence of lateral shoots that bear large amounts of fruit. Between
5 Bud Structure and Evolution 125

flower bud initiation and fruit maturity, it takes around 90–120 days. Pruning and
other techniques can maximize the production of flower buds and help timing the
production to market needs, but this practice should be done with care. Severe prun-
ing in ‘Col de Dame Noire’ and ‘Bourjasotte Noire’ caused a reduction in the aver-
age number of flower buds on shoots (Gerber et al., 2010). The flower buds of
brebas may wait until the season is somewhat advanced (i.e., warmer) before they
start active growth, depending on the world region. If the growth and ripening of
brebas are delayed, the tree may put much of its initial resources into vegetative
growth upon breaking dormancy. The use of plant growth regulators (PGRs) and
other means to control bud burst can be an important factor. Cyanamide salts, such
as calcium cyanamide, are PGRs that are effective regulators of flower buds (bre-
bas) break (Shulman et al., 1986; Yablowitz et al., 1998). Applying 1–5% cyana-
mide advances bud break by 14–20 days compared with non-treated trees.
Applications of 1–5% cyanamide with ethylene (400 ppm) further advance bud
break by 20–30 days compared to non-treated trees. In addition to, or as a replace-
ment for cyanamide treatment, GA (gibberellic acid) can be applied to stimulate the
breba fruit and temporarily slow vegetative growth of the terminal mixed bud. Once
a terminal bud begins active vegetative growth, it suppresses (or slows) those breba
fruits located on the 1-year-old wood. However, if the tree is sprayed with gibberel-
lins (for example, GA3 at a concentration of about 40–50 ppm), usually two times at
1-week interval, the growth of brebas will be significantly stimulated. The optimal
time for application is as soon as swelling of the terminal mixed bud is observed.
TDZ (N-phenyl-N-1,2,3,-thidiazol-5-ylurea), a cytokinin analogue, and mineral
oil have been used as an exogenous application, especially in apple, to overcome
dormancy (Steffens & Stutte, 1989; Wang et al., 1991; Honeyborne, 1996).
Girdling consists of removing a ring of bark (phloem tissue) from the shoot or
branch to restrict the movement of assimilates from the aerial portion of the tree to
the basal portion (trunk and roots), so the sap flow (mainly carbohydrates but also
hormones) is directed to buds, flowers, and fruits above the girdled zone. The inter-
ruption of phloem vessels is temporary (several days), and the tree then restores the
vessels through the production of a callus, a mass of parenchymatic cells (Davie
et al., 1995; Theron & Steyn, 2008; Ferrara et al., 2014). Interruption of the auxin
transport causes a sharp rise in cytokinin content leading to a stimulation of the
meristematic activity, which could cause an increase in flower bud induction and
initiation (Theron & Steyn, 2008).
Notching involves cutting through or removing a strip of bark directly above a
bud, effectively disrupting transport in the phloem. Notching involves an interrup-
tion of the influx of auxins which do not exert their inhibiting action and allow the
bud to develop (Greene & Autio, 1994). Notching was successfully used in the fig
‘Brown Turkey’ to increase bud break (Valia et al., 1994). When main management
factors (i.e., irrigation, fertilization) are in a proper balance, the new shoots will
continually produce new flower buds, leaves, and fruits. Overabundance of nitrogen
fertilizers can accelerate vegetative growth during the season but delaying the devel-
opment of the flower buds.
126 G. Ferrara and A. Mazzeo

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Chapter 6
Phenotypic Variability of Fig (Ficus
carica L.)

Ali Khadivi and Farhad Mirheidari

1 Introduction

Fig (Ficus carica L.) is one of the oldest domesticated fruit species in the world. The
English word fig is of ancient origin, derived from the Latin ficus. The species name
carica refers to Caria, an ancient region of Asia Minor noted for figs (Ferguson
et al., 1990). The genus Ficus is one of 40 genera belonging to the Moraceae family
with more than 800 species and most found in the tropics or subtropics, the most
important of which are F. carica, F. religiosa, F. elastica, F. benghalensis, and
F. rumphi (Condit, 1969; Woodland, 1997; Berg & Corner, 2005).
The genus Ficus is found mainly in the tropics, but Ficus carica L. is a subtropi-
cal fruit tree (Ferguson et al., 1990). Figs are currently cultivated in warm and tem-
perate climates and have grown as a Mediterranean fruit tree for many years (Sahin,
1998). The ancestral fig (Ficus carica L.) tree was monoecious and later evolved
into gynodioecious species with bisexual trees (functional male figs or caprifigs)
and unisexual female trees (Nabli, 1989; Machado et al., 2001). Both female and
male flowers are organized inside the syconium. A female fruit contains hundreds of
female flowers. A male fig plant, called caprifig, has hundreds of separated male
(staminate) flowers and female (pistillate) flowers. The differentiation between the
hermaphroditic and female strains forms establishes the basis for maintaining the
close symbiotic relationship between Ficus plants and the Blastophaga wasps (Galil
& Neeman, 1977).
The common fig tree is indigenous to Persia and Syria (Ferguson et al., 1990).
The fig tree has been distributed from Persia, Asia Minor, and Syria by people

A. Khadivi (*) · F. Mirheidari

Department of Horticultural Sciences, Faculty of Agriculture and Natural Resources,
Arak University, Arak, Iran
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 129
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_6
130 A. Khadivi and F. Mirheidari

throughout the Mediterranean area. Fig is considered the oldest cultivated crop
(Storey, 1975; Kislev et al., 2006). Fig cultivation has been related to its pleasant
taste and medicinal properties (Khadivi et al., 2018). Thousands of cultivars, pri-
marily unnamed, have been developed or came into existence as human migration
brought the fig to many places outside its natural range. They are now grown in most
Mediterranean countries, Europe, North and South Africa, the Americas, and many
other Asian countries (Flaishman et al., 2017).
Variability within fig accessions is attributed to the wide genetic diversity of fig
germplasm, as revealed by studies based on morphometric parameters (Mars et al.,
1998; Caliskan & Polat, 2008; Gaaliche et al., 2012; Khadivi et al., 2018; Mirheidari
et al., 2020; Hssaini et al., 2020b) and molecular markers (Khadari et al., 1995a;
Aksoy et al., 2003; Stover & Aradhya, 2008; Baraket et al., 2009; Simsek & Yildirim,
2010; Podgornik et al., 2010; Giraldo et al., 2010; Almajali et al., 2012; Simsek
et al., 2017). In addition, researching various genotypes of wild edible figs will
allow the fruit growers and breeders to select the most exciting and appropriate
range of wild edible fig accessions based on market needs taking into account the
ripening season and fruit color (Mirheidari et al., 2020; Khadivi & Mirheidari,
2022). Furthermore, the cultivated fig cultivars’ performance varies under different
climatic conditions (Ge & Zhang, 2006; Podgornik et al., 2010; Jacob et al., 2014;
Gruda & Tanny, 2015; Saadi et al., 2015; Caliskan et al., 2017). Cultivars also vary
in susceptibility to diseases (Yakushiji et al., 2012), with yield quantity and fruit
quality depending enormously on cultivation practices and climatic conditions
(Gaaliche et al., 2011).
A crucial genetic erosion occurs due to biotic and abiotic processes (urbaniza-
tion, extension of intensive crops, absence of caprification, water deficit in marginal
areas, etc.) (Mars et al., 1998; Salhi-Hannachi et al., 2004). To better conserve and
use genetic resources, characterization designs of morphological variability within
the collections and selecting the most significant variables to establish strategies for
better conservation and use of genetic resources should be performed carefully
(Giraldo et al., 2010). Despite the advances in molecular characterization in fig
(Achtak et al., 2009; Giraldo et al., 2005, 2008a; Ikegami et al., 2009; Khadari et al.,
2005), phenotypic characterization is always needed, and it should be included in
any program of conservation and use of genetic resources (Giraldo et al., 2008b).
Ideally, this characterization should be made under the same edaphoclimatic
conditions.

2 Botany and Biology in Different Types of Figs

Most species of the genus Ficus are monoecious and have male and female flowers
within single syconia. Ficus carica is a gynodioecious woody perennial species
with two tree types: male fig trees (the inedible caprifig) produce male flowers and
short-style female flowers, while female fig trees (the edible fig trees, i.e., Smyrna,
San Pedro, and common type) only produce long-style female flowers (Lisci &
6 Phenotypic Variability of Fig (Ficus carica L.) 131

Fig. 6.1 (a) Monoecious caprifig fruit and (b) Female fig fruit. Blue, red, and green arrowheads
indicate staminate flower, short-style pistillate flower, and long-style pistillate flower. (c) Caprifig
staminate flowers (left), caprifig short-style pistillate flowers (center), and fig long-style pistillate
flower (right) (Mori et al., 2017)

Pacini, 1994; Valdeyron & Lloyd, 1979; Stover et al., 2007a) (Fig. 6.1). Although,
some so-called caprifigs are reported to be edible and have a more succulent fruitlet
than typical caprifigs (Storey, 1975).
Caprifigs (male figs) of Ficus carica var. caprificus Risso produce three crops of
syconia per year: the profichi that ripen in early summer, the pollen source for the
edible fig, are produced in large numbers on wood from the previous season; the
mammoni that ripen in fall; and the mamme that over-winter on the tree and ripen in
spring. Fig pollen is transferred from male flowers to female flowers by an insect
known as the fig wasp (Blastophaga psenes, Hymenoptera: Chalcidoidea:
Agaonidae); they represent perhaps the most extreme and ancient (~75 million
years) obligate pollination mutualisms known (Machado et al., 2001; Cruaud et al.,
2012). These wasps live, lay their eggs, and feed within the fig syconia. Only the
winged female wasps emerge and leave the fig through the ostiole (wingless males
cannot leave the syconia). The female wasps fly to new syconia to oviposit eggs in
some female flowers, one egg per ovary. However, the profichi caprifig has many
male flowers near the ostiole, and the pollen from these flowers is carried by the
female wasps to the next syconium, pollinating the ovaries. Although caprifig
132 A. Khadivi and F. Mirheidari

syconia incubate and perpetuate these tiny fig wasps, seeds may also develop in the
female flowers. This is especially true of mammoni syconia in which pollination
results in some ovaries (without wasps) developing seeds with viable embryos
(Condit, 1955; Galil, 1977).
Three different types of female figs could be distinguished depending on the
cropping/pollination characteristics, common type, Smyrna, and San Pedro (Stover
et al., 2007a). Most fig genotypes are common (e.g., ‘Brown Turkey’, ‘Mission’,
‘Adriatic’, Dottato, etc.) that produce parthenocarpic fruit without caprification. The
drupelets inside develop parthenocarpically, and the endocarps are generally hollow
without fertile seeds. Common type figs can produce one (unifera types) or two
crops (bifera types). The Smyrna types (e.g., ‘Calimyrna’ (‘Sarilop’), ‘Black Bursa’,
‘Zidi’, ‘Sabz’, etc.) require pollination with pollen from caprifigs (caprification) for
fruit development to occur. These trees produce only two crops of syconia annually,
a breba crop that ripens in early summer and the main crop that ripens in autumn.
Finally, the San Pedro types (e.g., ‘Dauphine’, ‘King’, ‘San Pedro’, ‘Lampiera’,
‘Petrelli’, etc.) can produce two crops; the first crop (breba) is parthenocarpic (it
proceeds directly without pollination and fecundation) and the second crop, like the
Smyrna, is produced only after caprification (Condit, 1955; Galil, 1977; Flaishman
et al., 2008a). This complex reproductive biology allows us to identify some of the
botanical features of figs and use them to distinguish more precisely between differ-
ent types of figs (Table 6.1).
The last two types of edible figs are not parthenocarpic (persistent) and require
pollination to set the main crop of figs. Botanists use the term “persistent” rather
than “parthenocarpic” because the fig is not a true fruit. Botanically, these nonper-
sistent types are classified as “caducous”. Of the cultivars described by Condit
(1955), 78% are common types, less than 4% are San Pedro types, and the remain-
ing 18% are Smyrna types.
In some varieties, the main crop could be divided into two sub-groups, the (sum-
mer) main crop (ripening in July-September) and the late main crop ripening in
autumn and borne on the trees up to December. Some varieties produce only the
main crop ripening late (cv. Natalino) and are eaten almost at Christmas (Marcotuli
et al., 2020). In the two-crop variety, breba fruits are considerably heavier and have
a shorter ripening time than the main crop, but in some years, there are not present
(Podgornik et al., 2010). However, this difference in crops leads to the distinction of
the varieties in uniferous (only one crop, main crop), biferous (two crops, breba and
main), and even triferous (breba, summer, and late main crop) (Ferrara et al., 2017;
Marcotuli et al., 2019).

3 Germplasm Diversity and Conservation

The increasing concerns about reduced levels of genetic diversity in crop species
(Esquinas-Alcazar, 2005; Tanksley & McCouch, 1997) have led to the need to pre-
serve as much genetic diversity as possible both in situ and ex situ not only for the
6 Phenotypic Variability of Fig (Ficus carica L.) 133

Table 6.1 Fig types and sexual characteristics (Bearing time can slightly vary in different
ecological areas)
Female figs
Male figs Caducous
Fig types Caprifig Smyrna San Pedro Common type
Flower Male flower with Male flower not Male flower not Male flower not
morphology five stamens, presented, female presented, female presented,
female flower with flower with long flower with long female flower
short style style style with long style
Presence of Male and female Only female wasp, Only female wasp, No wasps
wasp wasp (wasps’ eggs from a caprifig from a caprifig involved
can develop only in syconia (wasp syconia (in the
short style flowers) cannot lay its eggs mammoni crop)
in long style
flowers)
Mamme (in Not fecundation Not developed Not developed Not developed
male figs) (pollen is not
(autumn- viable), small fig
winter) not edible
Profichi (in Production of Fecundation by Not fecundation
Not
male figs) pollen in the male pollen from a (persistent), setting
fecundation
Breba (in flower but no carprifig, edible on wood from the
(persistent),
female figs) fecundation, fig not fig, but it cannot previous year,setting on
(spring- edible be dried edible fig, but it
wood from the
beginning cannot be dried.
previous year,
summer) edible fig, but it
cannot be dried
(developed
only in bifera
types)
Mammoni (in Fecundation by Fecundation by Fecundation by Not
male figs) pollen from profichi pollen from a pollen from a fecundation
Main crop (in (mammoni does not caprifig, edible fig carprifig, setting on (persistent),
female figs) produce pollen), and it can be dried current season’s setting on
(summer- small and hardly wood, edible fig, current
autumn) edible fig can be dried. season’s wood,
edible fig and it
can be dried
(developed in
both unifera
and bifera
types)
Seed Fertile seed Fertile seed Infertile seed Infertile seed
characteristic (embryo and (embryo and profichi (hollow (hollow seed)
endosperm endosperm seed), fertile seed
developed) developed) mammoni (embryo
and endosperm
developed)
Adapted from Mateos and Gil (2014)
134 A. Khadivi and F. Mirheidari

long term survival of the species but also to have enough variability available for
breeding programs. Although several efforts have been carried out to conserve fig
genetic resources in germplasm banks (Flaishman et al., 2008b; Giraldo et al.,
2008a; Stover & Aradhya, 2008), appropriate management of the germplasm col-
lections requires defining patterns of phenotypic variability within the collections
and selecting the most significant variables to establish strategies for better conser-
vation and use of genetic resources. For this purpose, the analysis of genetic diver-
sity in germplasm collections facilitates reliable classification of the accessions
conserved and the identification of subsets or core accessions with possible use for
specific breeding purposes (Mohammadi & Prasanna, 2003; Escribano et al., 2008).
Therefore, it is imperative to establish programs to preserve and characterize fig
genetic resources globally, and programs have been developed in several countries.
Fig germplasm accessions are reported in Asia, Europe, North Africa, and North
America. The largest collections are in Italy, Turkey (Elbreyli Fig Research
Institute), California (National Clonal Germplasm Repository (NCGR) Davis),
Turkmenistan (TES-PGR), and France (CBNM Porquerolles), containing 442, 314,
190, 180 and 154 accessions, respectively. The CBNM Porquerolles collection has
154 different fig cultivars from Italy, North Africa, Greece, Turkey, France, Tunisia,
and Spain. The NCGR fig collection contains 190 accessions. The Arid Land and
Oasis Cropping Laboratory germplasm collection of the Institute of Arid Regions of
Medenine, Tunisia, holds 119 local fig accessions and 20 caprifigs collected from
Tunisia, Algeria, and Egypt (Aljane et al., 2012).
To better conserve and utilize genetic resources, characterization designs of mor-
phological variability within the collections and selection of the most significant
variables should be carefully performed (Giraldo et al., 2010). This characterization
usually involves a wide range of data which include both qualitative and quantita-
tive traits (Khadivi-Khub et al., 2012). It is worth noting that some of these param-
eters are generally influenced by environmental conditions and agronomic practices.
Nevertheless, morphological characterization is a highly recommended first step
before starting biochemical or molecular studies (Hoogendijk & Williams, 2001).
Several studies have reported the morphological characterization of female fig
cultivars and proved the sustainability of the use of morphological parameters to
evaluate and establish a description of genotypes (Condit, 1941; Mars et al., 1998;
Oukabli et al., 2002; Hedfi et al., 2003; Salhi-Hannachi et al., 2003; Aljane et al.,
2012; Gaaliche et al., 2012; Caliskan & Polat, 2008, 2012; Almajali et al., 2012;
Khadivi et al., 2018; Rodrigues et al., 2019a; Mirheidari et al., 2020; Hssaini et al.,
2020b, c; Moniruzzaman et al., 2020; Khadivi & Mirheidari, 2022).
Authors stress the relevance of morphoagronomic variability in cultivars’ identi-
fication and breeding programs and claim such analysis should be performed before
molecular studies are carried out (Podgornik et al., 2010; Khadivi-Khub et al., 2012;
Djordjević et al., 2014; Khadivi et al., 2018).
6 Phenotypic Variability of Fig (Ficus carica L.) 135

4 Wild Figs, Cultivars, Local Clones, and Hybrids

4.1 Wild Fig

Fig (Ficus carica L.) is a traditional fruit from West Asia and the eastern portion of
the Mediterranean, including Turkey and Iran, in which wild variants of this species
are found (Ikegami et al., 2009). These wild species consist of ecotypes called com-
mon figs (unisexual female trees) and caprifigs (bisexual with functional male trees)
occurring in similar frequencies in wild populations (Valdeyron & Lloyd, 1979). In
Tunisia, the wild fig is mainly found in the north, and the center, such as in the
islands of Galite and Kerkennah, Djebba, El Haouaria, and Kesra and is widely
spread over all the climatic stages (Mars et al., 1998; Aljane et al., 2005). Wild fig
is seed-propagated, and seeds are disseminated by birds (Lisci & Pacini, 1994).
Therefore, it is primarily present in rocks, usually along the riverbanks, in steep-
sloped valleys (Khadari et al., 2005).

4.2 Cultivars and Local Clones

Many of the existing fig tree cultivars may result from selection focusing on agro-
nomic characteristics and/or selection and transportation to distant regions from
breeders and growers (Condit, 1955). Stem-cuttings usually carry out fig propaga-
tion, and this procedure has contributed to synonymy and homonymy because mis-
identification and somaclonal variations are common events in sexually propagated
species such as fig (do Val et al., 2013). The long period of domestication, the use-
fulness of the perpetuation of certain traits through asexual propagation, and the
exchange of materials also contributed to synonymy (Aradhya et al., 2010).
Fig cultivars are numerous, and genotypes are either misidentified or called by
different denominations in different regions of the world (Aljane, 2016; Aljane
et al., 2012; Essid et al., 2017). Fig names were mainly given based on phenotypic
and morphological characters such as fruit color, fruit shape (width and neck
length), fruit taste and flavor, local geography, and maturity period (IPGRI and
CIHEAM, 2003; Caliskan & Polat, 2008; Aljane & Ferchichi, 2009; Hssaini et al.,
2019). For example, the variety ‘Ghoudan’ indicates the darkness of skin color, and
‘Zerqui’ refers to the blueness of fruit skin. ‘Ounq Hmam’ refers to the length of the
neck (long fruit neck) and means in the local dialect “pigeon neck.” Denomination
of ‘Jaadi’ describes genotypes having ribs over the skin. ‘Nabout’ is a generic term
that means “tree that grows spontaneously” (Hmimsa et al., 2012). Moreover, the
same variety may have different names depending on the cultivated region. For
example, the cultivar ‘Sari-Lop’ (‘Smyrna’) from Asia referred to California as
‘Calimyrna’.
Regarding the phenotypic diversity between fig cultivars, Condit (1955) has
already provided a comprehensive list of fig varieties, and recently the names and
136 A. Khadivi and F. Mirheidari

characteristics of some cultivated accessions/cultivars in different countries have

been reviewed (Singh et al., 2015; Hiwale, 2015; Núñez-Gómez et al., 2021).
In Turkey, fig culture is seen in almost every part of the country except the very
continental climatic areas of the Middle and East Anatolia. Almost all the coastal
lines of the country, such as the Aegean, Mediterranean, Marmara, and Black Sea
regions, are the main areas for fig production (Arpaci, 2017).
Most Turkish dried and fresh fig cultivars, ‘Sarilop’ and ‘Bursa Siyahi’ are cadu-
cous and classified as Smyrna types, respectively (Ozen et al., 2007; Arpaci, 2017).
‘Bursa Siyahi’ is Turkey’s most commonly grown fig cultivar (Aksoy et al., 2003).
Fruit characteristics of the fig cultivars grown in Antalya conditions (Bursa Siyahi,
Yesilguz, Bardakci, Sultan Selim, Karabakunva, Beyaz Orak, and Sarilop) were
studied by Kaynak et al. (1998). Bursa Siyahi had the heaviest fruits and Karabakunya
the lightest. Fruit width was greatest in Beyaz Orak, Sarilop, and Bursa Siyahi, and
Bursa Siyahi had the longest fruits. Bursa Siyahi, Sarilop, and Yesilguz were the
easiest to peel. Bardakci, Bevaz Orak, Bursa Siyahi and Yesilguz had the least osti-
ole and skin cracking. Also, five local cultivars (Degirmen, Ipek, Istanbul, Pamuk,
and Pathcan) in Northern Turkey are predominantly consumed as fresh fruit. In
addition, some of these cultivars are used for making jams. The harvest period is
long, from early September to late October. Therefore, single cropping systems of
large fig plantations are essential in optimizing economic production (Bostan
et al., 1998).
El-Kassas et al. (1992a) evaluated the growth of four local and introduced culti-
vars in Egypt. The cultivars DiRedo, Conadria, Sultani, and Black Mission, were
best in terms of tree vigour. However, the yield and quality of syconium also showed
marked variation in different cultivars (El-Kassas et al., 1992b).
The most common cultivars grown in Morocco are ‘Nabout’, ‘Ghoudan’,
‘El-quoti lbied’, ‘Ghani’, ‘Chaari’, ‘Fassi’, and ‘Ounq Hmam’ (Hssaini et al.,
2020b). The Riffian Mountains in Northern Morocco are the focal area of fig plant-
ing and production, which is considered a source of income for the local farmers
(Hmimsa et al., 2012). Also, Hssaini et al. (2019) indicated that the most dominant
varieties in Northern Morocco (Taounate, Ouzzane, and Meknes regions) were:
‘Nabout’, ‘Ghoudan’, ‘El Quouti lbied’, ‘Zerqui,’ ‘Ghani’, ‘Sbeti’, ‘Tabli’, ‘Lamtal’,
‘Mssari’, ‘Ounq Hmam’, ‘Arguil’, ‘Chaari’, and ‘Jaadi’. According to their survey
results, these varieties were all propagated using hardwood stem cuttings. In addi-
tion, Hmimsa et al. (2012) counted about 133 denominations in the Rif Mountains
in Northern Morocco. Furthermore, they reported 191 lexemes corresponding to
133 morphotypes or varieties reproduced by vegetative propagation in the Rif area.
Furthermore, they concluded that names might vary from one locality to another for
the same morphotype, which explains the problem of mislabeling that characterized
the cultivated varieties in the Rif mountains.
Iran is a favorable country for fig growing. In Iran, fig culture is seen in almost
every country. It is also found in the wild type (edible fig and caprifig) in most parts
of the country. Dry farming of figs in the Fars province of Iran started about
650–700 years ago, and most of the land cultivated with dry-farmed figs is located
in Fars province (Rahemi & Jafari, 2008). The Smyrna-type cultivars represent the
6 Phenotypic Variability of Fig (Ficus carica L.) 137

main type commercially grown in Iran. Estahaban is the main region of fig fruit
production in Iran (Khadivi et al., 2018). The fig landrace ’Sabz’, the main dried fig
in Iran, belongs to the Smyrna type figs that bear only pistillate flowers and require
the wasp pollinator to pollinate its fruit. The fruit crop of ’Sabz’ ripens and partially
dries on the tree. The’Sabz’ landrace fruit crops are dried directly on the tree and do
not fall until perfectly ripened. Instead, they are gathered after fallen and left in the
sun for a few hours before being conditioned for sale (Khadivi et al., 2018). Also,
other commercial cultivars include ‘Payves’, ‘Sigoto’, ‘Khafrak’, ‘Rowno’,
‘Shahanjir’, ‘Atabaki’, ‘Kashki’, ‘Matti’, ‘Manbili’, and ‘Seyah’ in Fars province in
the south-west (Pourghayoumi et al., 2017; Khadivi et al., 2018) and ‘Lashei’,
‘Majifi’, ‘Bar-anjir’, ‘Sham’, ‘Bavameli’, ‘Shamamleh’, ‘Zardleh’, ‘Siavleh’,
‘Estahban-e-Sabz’ in Kermanshah and Ilam province in the west (Fatahi et al.,
2017), ‘Siah Esfehan’, ‘Emadi Sabze Esfahan’, ‘Poost sabze eghda’, ‘Peyvandiye
Varamin’, ‘Siyahe Mahali’, ‘Zard Bongah Varamin’ in the center and ‘Peykami
kashmar’, ‘Zarde Boshghabi Kashmar’ in east of Iran (Mahdavian et al., 2008).
In 1850, more than 70 fig varieties were introduced to Algeria. However, these
were not adopted by the local farmers, who continued to grow the fig trees that were
familiar to them (Mahmoudi et al., 2018). The Technical Institute of Fruit-bearing
Arboriculture in Algeria described 40 varieties, including comestible and caprifig
types (Chouaki, 2006; Meziant et al., 2015). The cultivar ‘Kalamon’ is the main
variety cultivated to produce dried figs in southern Peloponnese and is represented
in FTC-IK by three accessions originating from Messinia. On the other hand,
‘Kalamatiani Evias’ that represents another accession cultivated in different parts of
Greece for the same commercial purpose, shares identical morphological character-
istics with ‘Kalamon’ and is, therefore, believed to be the same cultivar (i.e.,
‘Kalamon’) (Papadopoulou et al., 2002).
Prospection undertaken in different regions in Tunisia contributed to identifying
and describing numerous cultivars (Ben Salah & Lejri, 1995; Mars et al., 1998;
Chatti et al., 2004; Mars et al., 2009). Some, like ‘Bither’ and ‘Bither besbessi’, are
of the common type (parthenocarpic) that produce figs without caprification (polli-
nation). Many others, like ‘Bidhi’, ‘Zidi’, ‘Kahli’ and ‘Soltani’, are Smyrna types
that need caprification. Smyrna types are predominant in the south while, in the
north, cultivars of common type are equally represented (Mars, 2003). The fig
germplasm consists of many ecotypes selected for their fruit traits (‘Bidhi’ and
‘Zidi’) or their high adaptative potentialities (‘Bouhouli’ and ‘Wahchi’). The culti-
var Zidi (Smyrna type) has a large geographical distribution in the country and is
known for its high commercial value and best fruit taste for fresh consumption
(Mars et al., 2008). Mars et al. (1998) analyzed the diversity among 22 cultivars of
the southern arid regions in Tunisia using morphometric studies. Cultivar character-
ization and multivariate analysis based on 18 physical and chemical fruit characters
of 22 main crop varieties suggested that fig germplasm is diverse. It was also pos-
sible to differentiate three cultivar groups and two distinct cultivars.
In Spain, almond and fig tree cultivation coexist with grape cultivation, intro-
duced during the twentieth century by immigrant farmers from the ‘Levante’ region,
eastern Spain (Perez-Jiménez et al., 2012). The main fig fruit-producing areas in
138 A. Khadivi and F. Mirheidari

Spain are Extremadura-Andalusia and neighbouring regions, the Levantine area,

and Galicia (Pérez-Sánchez et al., 2016). Most of the varieties cultivated in Spain
(e.g., ‘Colar’, ‘Super-Fig’, ‘Cuello de Dama Negro’ and ‘San Antonio’) belong to
the parthenocarpic and biferous group, but there are also those of the San Pedro type
(i.e., ‘Nazaret’ and ‘Tiberio’ varieties) mainly cultivated in Spanish western (López-
Corrales et al., 2011; Núñez-Gómez et al., 2021). In addition, ‘Carballar Negra’ and
‘Moscatel’ are the only cultivars with one crop per year (unifera type) (Pérez-
Sánchez et al., 2016).
Despite its socioeconomic and historical importance, in Italy, the fig is consid-
ered a minor fruit species. The regions in which the fig tree assumes economic
importance are Campania (Cilento-SA) and Calabria (province of Cosenza), fol-
lowed by Puglia (Salento area-LE) and Sicily (Ciarmiello et al., 2015). In Calabria
and Campania, the main production of dried figs is based on the cultivar ‘Dottato’,
admitted in two different protected designation of origin (PDO): “Fichi di Cosenza”
(Calabria) and “Fico Bianco del Cilento” (Campania). The cultivar ‘Dottato’, syns.
= ‘Ottato’, ‘Uttato’, ‘Ortata’, ‘Fico Bianco’, ‘Kadota’, (Gallesio, 1820; Vallese,
1909; Condit, 1955; Grassi, 1991) is of ancient origin. ‘Dottato’ develops partheno-
carpically both the first crop (breba) and second-crop (figs), which represent the
main production.
The most famous Greek fig tree cultivar, ‘Kymis’, which is widely cultivated on
the island of Evia, has a central market position and high potential since it is in the
process to be appointed a Protected Designation of Origin (PDO) EU mark.
Currently, it is highly-priced, offering a significant income to local farmers (Vlachos,
2013). ‘Dauphine’ is another fig cultivar similar to ‘Masui Dauphine’, an important
Japanese cultivar popular in Japan. Masui Dauphine was evaluated for 5 years in the
Zhenjiang area in China. It produced large (80–100 g), elongate-oval, purple-red
fruits with 15% soluble solids content and high aspartic acid content (0.22 mg/100 g).
The fruit was suitable for fresh consumption and processing (Yang et al., 1994).
The main varieties cultivated in Cyprus are ’Vazanata’, ’Vasilika’, ’Smyrneika’,
’Vardika’, ’Kadota’, ’Napolitana negra’, ’Progontto Blanco’ and ’Gentile Bianco’
(Gregoriou, 1995). The main cultivars planted in California are “Calimyrna,”
“Mission,” “Brown Turkey,” and “Kadota” (Stover et al., 2007b). Most fig produc-
tion in California is used for the dried market; only a small percentage is harvested
for fresh fig consumption (Stover et al., 2007b). Also, Marseilles (White Marseilles,
or Lemon) is commonly found in California (Singh et al., 2015). ‘Black Mission’ is
cultivated in California on a large scale. This variety is old but highly productive. As
per the name, the fruits are black. The fruits are sweet and tasty and suitable for
drying and preparing jam. The first season’s fruits are bigger than the second season
(Hiwale, 2015). The Tree is a vast, elongated fruit, with Flesh watermelon to pink
and a reasonably good taste. Breba crop is prolific (Singh et al., 2015).
‘Adriatic fig’ trees could sometimes be found in the state of Washington. This is
a common variety in Chile and Argentina (Singh et al., 2015). ‘Celeste figs’ are
most commonly grown in Texas. ‘Brown Turkey’ is grown on all the islands
of Hawaii.
6 Phenotypic Variability of Fig (Ficus carica L.) 139

In India, fig farming is limited to a few states such as Maharashtra, Gujarat, Uttar
Pradesh, Karnataka, and Tamil Nadu. In northern India, its cultivation is spread to
some parts of Uttar Pradesh, Uttarakhand, Punjab, and Himachal Pradesh (Sharma
& Badiyala, 2006). ‘Poona’ and ‘Dinkar’ are widely acceptable varieties of fig.
Apart from these, the important varieties of figs cultivated commercially in India
are; Conadria, Deanna, Excel, Celeste, Brown Turkey, and Brunswick (Magnolia)
(Gani et al., 2018). Also, At Saharanpur, India, ‘Brown Turkey’, ‘Bangalore’, ‘Black
Ischia’ and ‘Lucknow’ are successfully grown. Around Mumbai, there is only one
variety, ‘Poona’ (Singh et al., 2015).
The fig tree has been cultivated in Brazil since the beginning of the last century,
and owing to the growing demand for these fruits by the food industry, the cultivated
area continues to increase each year, especially in the States of São Paulo, the Rio
Grande do Sul, and Minas Gerais (Paula et al., 2009). Fig cultivation in Brazil is
mainly based on a single cultivar, ‘Roxo de Valinhos’, introduced by Italian immi-
grants who settled in the State of São Paulo in the late nineteenth century (Paula
et al., 2009; Francisco et al., 2011). According to Ferreira et al. (2009), the predomi-
nance of this cultivar in Brazilian orchards is due to its desired traits of vigor, pro-
ductivity, rusticity, and wide acceptance by consumers and the industry.
In Costa Rica, the main fig cultivar produced is ‘Brown Turkey’, and it has been
being found in Tierra Blanca and Pacayas (in the north region of the Cartago prov-
ince) since the colonial period as a complementary agricultural activity to the potato
and onion crops (Flores et al., 2011).

4.3 Hybrids and Transgenic Varieties

The classical breeding approaches in fig were summarized by Condit (1947), Storey
(1975), Obenauf et al. (1978), Ferguson et al. (1990), and Mars (2003). However, fig
breeding is complex because of involving two tree morphs (Caprifig and edible fig),
three floral forms (long-styled female, short-styled female, and male flowers), and
the insect pollinator (Beck & Lord, 1988). These cause problems in breeding proce-
dures like; the occurrence of parthenocarpy and seedlessness, complex pollination
mechanism, i.e., caprification, lack of reference collections, lack of universal
descriptor list to differentiate different varieties/species, presence of large intra-
variety diversity, the closeness of wild types to cultivated plants in some regions
(Beck & Lord 1988).
Fig crosses and their evaluation has been described in Japan (Awamiira et al.,
1996). In addition, two breeding programs were conducted at the beginning of the
twentieth century at the University of California. The most significant achievement
of the California fig breeding was the development of 5 hybrid cvs.; Conadria, Di
Redo, Flanders, Tena and Excel. Furthermore, a common cv. Sierra was developed
with late summer maturity and green color skin (Doyle & Ferguson, 2005).
Transgenic varieties are limited in fruit trees, and their efficiency is very diverse
according to the fruit species, some being recalcitrant (Chevreau, 2009).
140 A. Khadivi and F. Mirheidari

Nevertheless, transgenic varieties have been developed to improve agronomic traits,

fruit storability, and disease resistance. The transgenic approach for fig-tree breed-
ing was described by Yancheva et al. (2005), wherein an Agrobacterium-mediated
transformation of the cv. Brown Turley and Smyrna were realized. Technological
progress enabled gene transfer and permitted the production of desirable proteins in
fig fruits to enhance the nutritional and pharmaceutical constitution (Flaishman
et al. 2008b). Also, the objective of Ozen et al. (2017) research was to investigate
the segregation of leaf characteristics among ‘Sarilop’ (female parent) × ‘Kaba ilek’
(male parent) hybrids. The importance of their study was to acquire knowledge
about the inheritance of some leaf characteristics in Ficus carica. In conclusion,
except for central lobe length, leaf length, width, area, petiole length, and thickness
were lower than parents. This might indicate negative transgressive segregation in
the leaf characteristics mentioned in the ‘Sarilop’ × ‘Kaba ilek’ fig cross.

5 Morphological Characteristics

The characterization and evaluation of plant species consist of establishing an iden-

tity for each accession through the knowledge of a series of data aiming to describe
the different accessions of a collection of germplasm, using characteristics of inter-
est. Cruz et al. (2014) mentioned that the characters studied may be morphological,
physiological, cytological, biochemical, or molecular and should include attributes,
characteristics, or traits of a culture that are informative and useful for the charac-
terization and evaluation of specific types (i.e., accessions) of the studied species.
Currently, morphological markers are still important and widely used in different
fruit species, cultivars and genotypes and are appropriate for classification (Poledica
et al., 2012; Yilmaz et al., 2012; Koc & Bilgener, 2013).
Selecting highly discriminant descriptors is important to optimize the resources
necessary for a viable and reliable morphological characterization (Giraldo et al.,
2005). In addition, an adequate descriptor should allow the distinction between dif-
ferent accessions from the same culture to be feasible and valuable, avoid redun-
dancy, be environmentally stable, and be easily manipulated by the breeder (Grum
& Atieno, 2007).
Growers and breeders appreciate morphological traits such as fruit dimensions
and stalk shape and, nevertheless, good adaptation to the environment and good
commercial value (Trad et al., 2013b). Besides the fruit size and shape, ostiole
width, acidity, and TSS (Mars et al., 1998), plant and leaf characteristics such as
plant growth habit, plant vigor, number of leaf and lobe per shoot, leaf shape are
essential for genotype selection by grower and breeders (Papadopoulou et al., 2002).
The researchers who have suggested that fruit and leaf traits are essential factors
in differentiating and analyzing breeding materials, and addressing the morphologi-
cal characterization of fig cultivars, are Mars et al. (1998), Hedfi et al. (2003),
Saddoud et al. (2008b, 2011), Gaaliche et al. (2012), Aljane et al. (2012) and
Khadivi-Khub and Anjam (2014). Also, many studies (Condit, 1955; Mars et al.,
6 Phenotypic Variability of Fig (Ficus carica L.) 141

1998; Papadopoulou et al., 2002; Mars, 2003; Aljane & Ferchichi, 2007; Saddoud
et al., 2008b, 2011; Gaaliche et al., 2012; Almajali et al., 2012) revealed that mor-
phological traits are beneficial in identification and evaluation of fig germplasm by
estimating genetic relationships among fig genotypes.
Furthermore, high phenotypic variability was reported in female figs from Iran
(Mahdavian et al., 2006; Baziar et al., 2018; Khadivi et al., 2018; Mirheidari et al.,
2020), Tunisia (Mars et al., 1998; Chatti et al., 2003; Aljane et al., 2012; Gaaliche
et al., 2012; Essid et al., 2017), Turkey (Caliskan & Polat, 2008, 2012; Caliskan
et al., 2017), Morroco (Oukabli et al., 2002; Hssaini et al., 2019, 2020b), Spain
(Sanches et al., 2002; Pérez-Sánchez et al., 2016), Lebanon (Chalack et al., 2005),
Palestine (Ali-Shtayeh et al., 2014), Jordan (Almajali et al., 2012), and Slovenia
(Podgornik et al., 2010). These studies indicated that high diversity in pomological
and leaf-related traits could be used as an efficient marker system to discriminate
between fig genotypes (Khadivi et al., 2018).

5.1 Biological Characteristics

Biological characteristics include traits like crop setting fruit, pollination require-
ment for fruit set, date of terminal bud-burst (leafing) in Breba, main crop and late
crop, beginning of fruit maturation, Full maturity, harvest period, onset of caprifica-
tion, length of the caprification period, onset of leaf fall. Among them, crop setting
fruit (bifera, unifera, san pedro, smyrna, caprifig), pollination requirement for fruit
set in breba (caducous, persistent), and main crop (deciduous, persistent) are more
discriminated traits as discussed in the text. Other traits are unstable in different
climatic conditions and can change according to environmental factors, and they
can be used more for accession comparison in germplasm collections or a specific
climatic region.
One of the most important aims is to expand the sales time thanks to the cultiva-
tion of genotypes/cultivars with different ripening times (Khadivi et al., 2018). The
ripening time of the fruits in the 87 Iranian edible fig genotypes was variable and
ranged between very early and late. Ripening time was early in 66 genotypes, late
in 15 genotypes, very early in three genotypes, and intermediate in three genotypes
(Khadivi et al., 2018). Also, Mirheidari et al. (2020) reported that the ripening time
of Iranian wild edible figs ranged from early July to early September.
The time of fruits ripening for breba was evaluated by Ferrara et al. (2016) in
some Italian accessions; they divided the biotypes into three classes: early
(Columbro, Mariana and Fiorone nero; 10–15 June), medium-late (Fico del Vescovo
and Tauro; 20–30 June) and late (Arodio; 5–10 July).
In Lebanese fig accessions studied by (Chalak et al., 2008), maturity dates were
determined in comparison to Biadi accessions, which were considered as mid-
season (11–31 August) in all fig growth areas in Lebanon, despite their different
ecogeographic conditions and only one accession (American) was classified very
late (after 1 October). Also, in Slovenian fig accession studied by (Podgornik et al.,
142 A. Khadivi and F. Mirheidari

2010), the ripening period of all accessions lasted 5–15 days for the first crop and
15–60 days for the second crop. Breba fruits of most accessions mature in mid-late
June, except accessions ‘Zelena Bonka’, ‘Madona Rjava’ and ‘Madnici’, which
ripen in early July. These three accessions are mainly cultivated in the hinterland
zone. The second crop ripens in early August and continues to ripen into September.
Accessions with early second crop (ripening between 1st and 10th August) were:
‘Greka crna, ‘Belica-2’, ‘Belica-1”, ‘Belica-3’, ‘Kanora’, ‘Zuccherina’, and
‘Bjelica’, while late second crop (ripening between 1st and 30th September) is char-
acteristics to accessions ‘Kamberij-1’, ‘Madona Rjava’, ‘Zelena Bonka’, ‘Crnica’
and ‘Zimica’ (Podgornik et al., 2010).

5.2 Vegetative Characteristics

5.2.1 Growth Traits

The fig tree’s growth is seasonal and related to diverse climatic factors. This includes
a period of more growth where the vegetative and productive cycles are completed,
followed by a dormancy stage, influenced by the region’s low temperatures and the
length of the winter season (Flaishman et al., 2008a). The adaptation to the region
can be monitored through various indicators; for example, if there are adverse envi-
ronmental conditions, the growth of the trees will slow down (Rostami & Rahemi,
2013; Gholami et al., 2012). In addition, other factors can affect the growth and
development of the fig tree (such as the type of agronomical management, nutrient
availability, the soil conditions, the hydric balance, among others) (Melgarejo,
2000; Botti et al., 2001; Hallac Turk & Aksoy, 2011; Crisosto et al., 2015) which are
characteristic of a given location (Botti et al., 2001).
Tree growth habit, size of the tree, degree of branching, number of lobes per leaf
(Giraldo et al., 2010), the number and shape of lobes (Saddoud et al., 2008a), leaf
length, leaf width, leaf area, the density of hairs/spicules on the leaf’s upper surface,
and petiole thickness (Podgornik et al., 2010) were the vegetative traits used for the
discrimination of fig accessions.
Fig trees vary in their growth habits, ranging from open and drooping to upright
and compact (Ferguson et al., 1990). ‘Brown Turkey’ has a weeping willow type of
growth with compact, down-spreading branches, while the cultivars ‘Sierra’ and
‘Autumn Honey’ have a vigorous growth habit with upright-rising branches.
Individual trees in a favorable environment often reach a large size, while fig trees
in orchards are usually compact (Flaishman et al., 2008a). The most frequent tree
vigor observed by Khadivi et al. (2018) in Iranian fig cultivars was intermediate, and
the majority of the studied trees had an open and spreading habit. Also, the Shoot
color examined by Mirheidari et al. (2020) in Iranian wild edible figs was highly
variable and included white, light brown, brown, dark brown, gray-white, and gray.
In addition, in another study on Iranian figs by Darjazi (2011), Plant growth habits
were obtained spreading (for Zard, Siah bolbol riz), erect (for Paizeh, Siah diras),
6 Phenotypic Variability of Fig (Ficus carica L.) 143

and weeping habit (for Bidaneh, Siah zoodras, Morabaii, Hallavi riz and Hallavi
dourosht) and tree vigor was found high (for Morabaii), medium (for Bidaneh,
Paizeh, Siah zoodras, Hallavi riz and Hallavi dourosht) and low (for Zard, Siah
bolbol riz and Siah diras).
Podgornik et al. (2010) reported intermediate tree vigor and spreading habit for
Slovenian figs. Very diverse vegetative habits were observed by Pérez-Sánchez et al.
(2016) in traditional fig cultivars grown in central-western Spain, ranging from
semi-erect to weeping, and cultivars vigor ranged between weak (‘Carballar
Blanca’) and strong (‘Antigua’, ‘Blanca Común’ and ‘Prieto’). López-Corrales
et al. (2011) also observed considerable variability in the growth habit character of
Spanish fig cultivars.

5.2.2 Leaf Traits

Leaf biometric markers exhibit drastic shape variation within the same species,
reflecting both the heteroblastic development of the apical meristem from which
they are derived and the ontogeny of individual leaves as they expand allometrically
(Chitwood et al., 2016). Each aspect of leaf morphology provides different points of
view about its pattern. The number of lobes is a highly relevant factor for the dif-
ferentiation of “toothed” plants, as is also the case with the fig tree, which proved to
be a promising feature of selection of leaf morphology among the features evaluated
by Rodrigues et al. (2017), because it presents high heritability and high genetic
correlation, being little influenced by the environment.
Fig has typical bright green, single, alternate, and large leaves. Leaf characters
are quite stable and serve as an essential parameter in cultivar identification
(Ferguson et al., 1990). In addition, the traits related to leaves are important for the
assessment of landrace and taxonomic investigations (Papadopoulou et al., 2002).
Leaf characters include the number of leaves per shoot, leaf shape, lobes number
and shape, location of little lateral lobes, degree of leaf lobation/incision, shape of
leaf base, leaf length and width, leaf area, length of leaf stalk/length of leaf, leaf
margin dentation, leaf margin, density of hairs/spicules on leaf upper surface, den-
sity of hairs or spicules on lower surface, leaf color, petiole length, petiole thick-
ness, petiole cross-section, petiole color (Condit, 1947; IPGRI and CIHEAM,
2003), vein number, central and lateral lobe depth (Khadivi et al., 2018), leaf vena-
tion clarity, central lobe length, upper lateral vein length, and upper lateral lobe base
width (Mirheidari et al., 2020).
Different leaf shapes in fig trees, according to Condit (1947), are (1) Base calca-
rate, lobes linear, (2) Base cordate, five-lobed, lobes spatulate, (3) Base calcarate,
lobes lyrate, (4) Base calcarate, lobes latate, (5) Base cordate, three-lobed, (6) Base
truncate, (7) Base decurrent and (8) Leaf not lobed. In addition, according to IPGRI
and CIHEAM, 2003), various leaf base shapes (petiole sinus) include truncate, cor-
date, calcarate, decurrent, and open calcarate. Also, the shape of lobes in fig leaves
in different genotypes can be spatulate (narrower at the base and wider at the top),
linear (more slender and regular in shape), latate (wider lobes), or Lyrate (as in
144 A. Khadivi and F. Mirheidari

Ficus lyrata) and different leaf margin shapes in fig leaves are crenate, dentate, ser-
rate, double serrate and undulate.
In 92 Iranian wild edible figs, leaf shape showed substantial diversity among the
accessions, including base calcarate/lobes linear (13 accessions), base cordate/five-
lobed/lobes spatulate (11), base calcarate/lobes lyrate (5), base calcarate/lobes latate
(10), base cordate/three-lobed (12), base truncate (6), base decurrent (31), and leaf
not lobed (4). In addition, leaf base shape (petiole sinus) was highly variable and
included truncate (8 accessions), cordate (42), calcarate (30), and decurrent (12).
Leaf margin showed six forms, although the crenate form (54 accessions) was pre-
dominant. Petiole cross-section was round and flattened (57 and 35 accessions,
respectively) (Mirheidari et al., 2020). Also, Khadivi et al. (2018) found high vari-
abilities among 87 Iranian edible fig genotypes detected in the leaf’s traits such as
leaf dimensions, leaf lobe depth, and the leaf lobe number. Most of the genotypes
had five lobes per leaf. Central lobe depth ranged from 16.8 to 62.0 mm, while lat-
eral lobe depth ranged from 16.8 to 92.0 mm. Petiole length ranged from 16.8 to
58.0 mm, and petiole thickness ranged between 1.70 and 6.30 mm.
In 21 local cultivated fig accessions from Egypt and Libya studied by Mohamed
et al. (2017), leaf length ranged from 5.4–23.5 cm, while leaf width varied from
6–23 cm. Four different leaf edge shape in accessions was strait (e.g., Kahramany,
Abodey-Giza), waved (e.g., Sultani black, Sultani-Giza), zigzag (e.g., Hamouri,
San-Badr), and serrated (e.g., Barry, Mejahal). Also, leaf top shape showed three
forms, although round form (e.g., Bioudi, Aswany) was predominant. In their stud-
ied accessions, the number of leaf lobes ranged from one lobe (e.g., Green-yellow,
Sultani Red Siwa), three lobes (e.g., Mejahal, San-Badr), five lobes (e.g., Sultani
black, Sultani yellow), six lobes (e.g., Hamouri), seven lobes (e.g., Adsey-Giza,
Fayoumi), eight lobes (e.g., Black-Mission) and ten lobes (e.g., Aswany).
According to Medeiros (2002), the ‘Roxo-de- Valinhos’ cultivar presents large
leaves with five large and two smaller lobes; dark green color; compact texture,
somewhat rigid; creased margin; petiole sinus in the form of lyre and long petiole.
Rodrigues et al. (2019b) mentioned that the number of lobes was the most promis-
ing leaf morphological character among those fig tree genotypes examined in their
study due to its high heritability and high genetic correlation, corroborant to
Caliskan and Polat (2012), which used the number of lobes per leaf as one of the
essential parameters for the differentiation of fig genotypes in Turkey. The differ-
ences between Slovenian figs have been reported in the degrees of leaf lobation (the
central lobe length/the leaf length) (Podgornik et al., 2010).
Khadivi et al. (2018) studied edible Smyrna-type fig clones from the Estahban
region in Iran and observed the range of 62.20–138 mm and 41–153 mm for leaf
length and leaf width. Also, the highest values of fruit stalk length and fruit stalk
diameter were 20.10 mm and 2.95 mm, respectively. Also, in examined fig cultivars
from Varamin in Iran, the leaf blade length of the genotypes was changed between
12.00 and 26.50 cm, and leaf blade width differed between 8.25 and 18.60 cm
(Darjazi, 2011).
In Spanish figs cultivars studied by Pérez-Sánchez et al. (2016), leaf length was
varied from 24.7 to 36.3 cm, ‘Tardía Portuguesa’ being the cultivar with the longest
6 Phenotypic Variability of Fig (Ficus carica L.) 145

leaf. Its blade size was also significant, with a length of 29.4 cm and a width of
24.2 cm. Stalk length ranged from 4.89 (‘Moscatel’) to 13.3 cm (‘Carballar Negra’).
It was observed that the leaf blades were longer than the stalks in all the fig cultivars
(stalk length/blade length ratio: 0.22–0.65). Concerning the number of main lobes,
most cultivars showed three/five-lobed leaves. These results are consistent with
those reported by López and Guzmán (2007) and González and Grajal (2012) for
Spanish fig cultivars. ‘Carballar Blanca’, ‘Cuarterón’, and ‘Moscatel’ were the only
cultivars that showed entire leaves. The shape of central lobe varied between trian-
gular (‘Carballar Blanca’ and ‘Cuarterón’) and lyrate (‘Cuello de Dama Negro’).
López-Corrales et al. (2011) also observed a lyrate central lobe shape in leaves of
this latter cultivar. The shape of the leaf base was highly variable among cultivars
(truncate-strongly calcarate). Most cultivars did not have additional lobes regarding
the basal lateral lobes on the petiole sinus. They were only observed in the leaves of
the ‘Blanca Común’, ‘Carballar Negra’ and ‘Cuello de Dama Negro’ cultivars. The
most frequent petiole color was green. Other researchers who have addressed the
leaf morphological characterization of edible fig cultivars are Giaccone et al. (2017),
Baziar et al. (2018), Rodrigues et al. (2019a), Abdelsalam et al. (2019),
Shahinuzzaman et al. (2020).

5.3 Pomological Characteristics

Common fig cultivars are facultatively persistent and may produce persistent polli-
nated main-crop figs. Morphologically, the pollinated fig syconium creates true
fruits, while the non-pollinated fig syconium presents an enlarged inflorescence
with multiple long-styled pistillate flowers. ‘Autumn Honey’ and the ‘Brown
Turkey’ are two cultivars that produce caprified and non-pollinated figs on the cur-
rent year’s wood. The caprified ‘Autumn Honey’ fruits have darker purple skin color
and red pulp than the white, pink pulp of the non-caprified fruit. Usually, caprified
fruits are bigger and have longer storage ability (Rodov et al., 2005). Where
B. psenes wasps are present, caprified and non-caprified fruit may develop on the
same branch in common-type fig fruits (Flaishman et al., 2008a). In the case of the
‘Smyrna’ cultivar, its shape tends to change due to the caprification process indis-
pensable for the normal development of fruits of this pomological class (Aradhya
et al., 2010).
Traditionally, the amplitude and distribution of fig tree genetic diversity have
been widely assessed based on conventional pomological markers, even though the
expression of these markers is strongly affected by environmental and agronomic
conditions of the species (Caliskan et al., 2017; Rodolfi et al., 2018). Therefore, fig
assessment using morphological, pomological, and physicochemical traits is essen-
tial for the inventory, assessment, preservation, and breeding programs of genetic
resources (Podgornik et al., 2010). Also, Caliskan et al. (2017) and Khadivi et al.
(2018) have reported that the pomological characteristics are essential to evaluate
the variation in traits of edible fig accessions.
146 A. Khadivi and F. Mirheidari

The most important discriminators of fig fruits were the fruit weight, fruit length,
fruit diameter, fruit skin and flesh color (Saddoud et al., 2008a; Aljane & Ferchichi,
2009; Podgornik et al., 2010), fruit shape (Giraldo et al., 2010; Podgornik et al.,
2010), firmness of the fruit skin (Saddoud et al., 2008a; Podgornik et al., 2010), fruit
skin cracks (Saddoud et al., 2008a), production type, skin firmness (Giraldo et al.,
2010), fruit neck length (Aljane & Ferchichi, 2009; Podgornik et al., 2010), abscis-
sion of the stalk from the twig (Podgornik et al., 2010), stalk diameter, neck diam-
eter, ostiole diameter, ostiole opening, and flesh thickness (Aljane and
Ferchichi 2009).
Fruit Shape Different fruit shapes in fig trees, base on fruit index [(width/length)
= I], are mainly Oblong (I < 0.9), Globose (I = 0.9−1.1) and Oblate (I > 1.1). Also,
fruit shapes, according to the location of the maximum width, include; Ovoid (in the
middle), Bell-shaped (nearer to the neck), and Pyriform (nearer to the ostiole-end)
(IPGRI and CIHEAM, 2003).
Fig fruits can be vertically and horizontally symmetric or asymmetric shape.
However, symmetric fruits are more acceptable for consumers and more suitable for
packaging. Fruit index (width/length) is important in packing and transportation
(Condit, 1941). Also, the fig shape and its index are of great importance when it
comes to trade and is prized by consumers. Aljane et al. (2008) indicated that fruit
shape is important for packing and transportation. However, the globose shape is the
most suitable fruit shape (Condit, 1941; Caliskan & Polat, 2008) and a more domi-
nant character in fig accessions (Gozlekci, 2011; Caliskan & Polat, 2012, Mirheidari
et al., 2020; Hssaini et al., 2020b) and also, it is preferred for its suitability for pack-
aging and transportation (Condit, 1941; Benettayeb et al., 2017). Although, other
authors reported that oblate fruit shape is the more dominant character (Aljane
et al., 2008; Darjazi, 2011; Caliskan & Polat, 2012). The harvesting dates and grow-
ing areas can explain these contradictory reports.
Most of the analyzed cultivars had an oblate fruit shape among some Iranian
cultivars. Fruits of Siah bolbol riz and Morabaii were globose while those of Hallavi
riz were oblong. (Darjazi, 2011). Pérez-Sánchez et al. (2016) reported that the most
frequent fruit shapes in Spanish figs were turbinate or spherical-turbinate. ‘Cuello
de Dama Negro’ was the only cultivar that showed pyriform fruits in their study.
López-Corrales et al. (2011) observed that this cultivar has pyriform breba. The fruit
skin ground color was generally green-yellow, ‘Prieto’ (purple), ‘Cuarterón’
(purple-black), and ‘Cuello de Dama Negro’ (black) being the only exceptions.
Fruit flesh color ranged from amber to red. Finally, some relevant cultivars with
regard to breba quality were ‘Antigua’, ‘Cuarterón’, ‘Cuello de Dama Blanco’,
‘Gota de Miel’, ‘Pringo de Mel’ and ‘Tardía Portuguesa’. Their fruits were very
juicy and easily peeled. González and Grajal (2012) also observed different fruit
shapes with Spanish fig cultivars.
The morphological characterization of 45 accessions in Active Germplasm Bank
Fig tree accessions of the Faculty of Agricultural and Technological Sciences
(FCAT/UNESP) was studied by Rodrigues et al. (2019a). They reported that the
shape of accession fruits varied from elongated to oval. The fruit shapes for
6 Phenotypic Variability of Fig (Ficus carica L.) 147

‘Pingo-de-Mel’, ‘Irradiated Plant 440’, ‘Roxo-de-Valinhos’, and ‘White Genova’

accessions were oval, elongated, oblong pyriform, and flattened, respectively.
‘Roxo-de-Valinhos’ cultivar has a predominance of fruits with oblong-pyriform
shape, with a short and thick neck, almost without limit of separation with the body
from the receptacle, as described by Rigitano (1955).
Fruit apex shape was flat in 87 out of 92 accessions studied in Iranian wild edible
figs. Conversely, the fruit was symmetric in 83 out of 92 accessions investigated
(Mirheidari et al., 2020). Hssaini et al. (2019) studied local fig clones from Northern
Morocco and reported that most studied genotypes had the symmetric shape of
the fruit.
Fruit Weight, Width and Length Fig yield is influenced by several factors, includ-
ing cultivar, growth conditions, and harvest. Typical mature fig yield varies between
5 and 12 t/ha (Flaishman et al., 2008a). Sahin et al. (2001) explained that fig plants
produced optimum yields at 7 years. Besides other criteria, weight is the most
important fruit dimension, and fruit prices depend mainly on this (Pérez-Sánchez
et al., 2016). Previous studies of pomological assessment of figs demonstrated that
the most important characteristics affecting the commercial value of fruits for fresh
consumption are fruit weight (Aksoy et al., 1992), width, and length (Can et al.,
2001; Ateyyeh & Sadder, 2006; Podgornik et al., 2010). Also, Fruit dimensions are
of great importance in the packing and transportation of fig fruits (Condit, 1947).
Hence, small fruits are generally used for canning, whereas big ones are consumed
as fresh. Moreover, the fruit size reflects the tree’s proper maintenance (Tamboli
et al., 2015). However, these character is usually negatively influenced by the fruit
load on the tree. It is noteworthy that besides the genetic effect, fruit weight also
depends on the growing location, growing season conditions, agricultural practices,
as well as the interaction between the genotype and the maturity stage, which is
properly explained by the index of maturity (Gozlekci, 2011; Benettayeb et al., 2017).
In general, fruits developing along the shoot are larger at the lower nodes and
become smaller toward the top of the shoot. Selected cultural practices may exert a
positive impact on fruit size. For example, in ‘Black Mission’, cultivar girdling
increased fruit size, although late girdling produced less marked effects (Ferguson
et al., 2003).
Uniformity of the final fruit size in fig trees is very important in the sales market.
Particularly for fresh products, due to the short post-harvest life of the fig and its
perishability, the more uniform the product, the less time it takes to sort. On the
other hand, it does not reduce the price of a part of the product due to its smaller size.
In Moroccan figs studied by Hssaini et al. (2020b), fig height values ranged from
21.3 to 48.8 mm with an average of 34 mm. At the same time, the fruit width was
25.93–61.58 mm, with an average of 39.8 mm. These results agreed with previous
works (Gozlekci, 2011; Papadopoulou et al., 2002; Ferrara & Papa, 2003; Rodrigues
et al., 2018).
In wild Iranian fig accessions, fruit length varied from 11.2 to 34.1 mm, and fruit
width ranged from 14.1 to 39.3 mm. Also, Fruit fresh weight ranged between 0.56
and 28.69 g with an average of 8.33 (Mirheidari et al., 2020). Aljane and Ferchichi
148 A. Khadivi and F. Mirheidari

(2010) studied fig cultivars from Tunisia and reported that Mean fruit lengths varied
from 51 mm in cv. Wahchi to 80 mm in cv. Faouari and mean fruit diameter was
between 47 mm in cv. Soltani Abiadh and 76 mm in cv. Faouari
In Turkish fig accessions studied by Gozlekci (2011), fruit width was between
28.6 and 51.1 mm, and fruit length was between 28.3 and 54.7 mm. Khadivi et al.
(2018) studied edible Smyrna-type fig clones from the Estahban region in Iran and
reported a range of 1.86–7.15 g for dried fruit weight. Also, in their study, Dried
fruit length ranged from 19.8 to 36.2 mm, and dried fruit width ranged from 15.7 to
25.2 mm.
Darjazi (2011) reported that Fruit diameter ranged from 21 to 45 mm, and fruit
length was varied from 20 to 36 mm in fig cultivars from Varamin in Iran. Also, in
their study, the fresh fruit weight ranged from 8.0 to 43.5 g, and the ‘Paizeh’ cultivar
showed the highest fruit diameter and fresh fruit weight.
Among eleven local and introduced fig varieties in Morocco that were studied by
Hssaini et al. (2020a), the variety ‘Kadota’ had the heaviest fruit (61.5 g), followed
by ‘Breval Blanca’ and ‘Snowden’, which the average weights were, respectively,
46.4 and 44.0 g. Furthermore, the latter recorded the highest
Caliskan and Polat (2008) reported fruit weight ranged from 22.2 to 52.5 g on
many fig accessions sampled in the East Mediterranean region. Gozlekci (2011)
carried out a selection study on figs in the Kemer and Alanya districts, which belong
to Antalya provinces located west Mediterranean Region in Turkey, and found that
fruit weight was between 14.7 and 60.5 g in the Kemer district, while the fruit
weight of fig accessions from Alanya district varied from 13.8 to 48.5 g. Similar
work carried out on a large part of fig diversity in different countries showed wide
variability in fruit weights that varied from 30 to 90 g (Ozeker & Isfandiyaroglu,
1998), 24 to 92 g (Sahin et al., 2001), 28 to 107 g (Polat & Ozkaya, 2005), 12.3 to
99.4 g (Caliskan & Polat, 2008) and 12.3 to 99.4 g (Caliskan & Polat, 2012) in
Turkey, 24 to 58 g (Aljane et al., 2005), 24 to 58 g (Aljane et al., 2008) and 63 to
172 g (Aljane & Ferchichi, 2010) in Tunisia, 30 to 85 g (Podgornik et al., 2010) in
Slovenia, 16.4 to 77.7 g (Moniruzzaman et al., 2020) in Malaysia and 12.4 to 87.0 g
(Hssaini et al., 2020c) in Morocco.
Fruit Stalk (Shape, Cross-Section, Length and Width) and Fruit Neck (Length,
Width) Different shapes of the fruit stalk in fig trees are short and thick, enlarged,
and long and slender. Also, the fruit stalk cross-section includes Circular, flattened,
rhomboid, and triangular (IPGRI and CIHEAM, 2003).
In Iranian wild edible fig accessions studied by Mirheidari et al. (2020), fruit
stalk shapes ranged from variously enlarged to long and slender. Hssaini et al.
(2020b) studied local Moroccan figs and reported that most genotypes had a rounded
shape at the stalk.
Also, Darjazi (2011) reported that the fruit stalk was long and slender for three
cultivars (Siah bolbol riz, Zard, and Hallavi riz), and it was short and thick for the
remaining fig cultivars from Varamin province in Iran (Darjazi, 2011). In Iranian fig
cultivars studied by Khadivi et al. (2018), the highest values of fruit stalk length and
fruit stalk diameter were 20.1 mm and 2.95 mm, respectively.
6 Phenotypic Variability of Fig (Ficus carica L.) 149

A neck in figs facilitates picking the fruit from the tree and is thus associated
with easier harvesting (Trad et al., 2012). In addition, short neck length is reported
to cause fig fruit damage during harvest (Ozeker & Isfandiyaroglu, 1998; Gozlekci,
2011; Darjazi, 2011). In Iranian wild edible fig accessions studied by Mirheidari
et al. (2020), the highest value of fruit stalk length, fruit stalk width, and fruit neck
length was 34.97, 4.75, and 8.31 mm, respectively.
Among the evaluated Moroccan fig germplasm, the stalk length and width ranged
between 2.95 and 16.3 mm and 2.92 and 7.96 mm, respectively. The largest size was
recorded with ‘Mnedar 289’ and ‘Figue de Marseille’ followed by ‘Trojana’ and
then ‘El Ghani’. Neck Length ranged from 1.64 to 9.52 mm, and Neck Width ranged
from 2.22 to 15.6 mm. The longest fruit neck was recorded in Bioudi 2878 (9.52
mm), followed by Nardine (9.4 mm) and Kodota (9.2 mm) (Hssaini et al., 2020c).
In the same study by Hssaini et al. (2020a), ‘Fassi’ had the highest fig stalk
(10.58 ± 3.5 mm) while ‘kadota’ had the highest stalk width (6.59 mm). Furthermore,
the fruit neck was long in the cultivars ‘Kadota’ (9.17 mm) and ‘Palmeras’ (7.28 mm)
and short in ‘Fassi’ (2.29 mm), whereas it was absent in others such as ‘Ghoudan’,
‘Nabout’ and ‘Palmeras’. Also, in another study by Hssaini et al. (2020b), for the
neck length, the values ranged between 1.64 and 9.52 mm with an average of 7.01.
At the same time, neck width varied from 2.22 to 15.7 mm, with an average of
8.64 mm. The longest fruit neck was recorded respectively by “Bioudi 2878”
(9.52 mm), “Nardine” (9.4 mm), and “Kadota” (9.2 mm).
In Turkish fig accessions studied by Polat and Ozkaya (2005), fruit neck length
in fig accessions ranged between 1.0 and 8.9 mm. neck length was longest on
31-IN-15 (8.86 mm) and Sarilop (6.39 mm). Gozlekei (2011) also reported fruit
neck length between 1.97 and 17.4 mm among fig accessions from Turkey. In other
studies, the longest neck was 14.5 mm (Ilgın, 1995), 8.70 mm (Ozeker &
Isfandiyaroglu, 1998) and 8.01 mm (Polat & Ozkaya, 2005). The cultivar ‘Cuello
Dama Blanca’ was closed to ‘White Adriatic’, ‘Nabout’, and ‘Noukali’, since they
all had no fruit neck. They may be used for canning, whereas big ones like ‘Kadota’
may be recommended for fresh consumption.
Ostiole Width The ostiole is characterized as the opening for the entrance of the
Blastophaga psene wasp so that inflorescence pollination within the siconium can
occur to develop fig fruits (Lamas et al., 2009). The ostiole provides entry for fungal
decay and moisture loss during the maturity period. It also provides an entryway for
insects that vector diseases such as endosepsis that spread to healthy fruits
(Michailides & Morgan, 1998; Crisosto et al., 2011). Thus, a large ostiole in the fig
is an undesirable characteristic (Caliskan & Polat, 2008; Can, 1993). The smaller
ostiole width, the better fruit can be stored and protected from infectious agents
(Trad et al., 2012). In the specific case of the fig tree crop, an essential aspect of
determining the quality of fruits and their size is the small ostiole opening (Aljane
et al., 2013).
Cultivars with a large ostiole (Doster et al., 2002) are susceptible to ostiole split-
ting (Michailides, 2003) and are more susceptible to decay. The high susceptibility
of figs to decay and their fast ripening accelerates fruit softening (Çelikel & Karaçalı,
150 A. Khadivi and F. Mirheidari

1998), making fresh fig shelf life extremely short, lasting only 1–2 days
(Morton, 2000).
In Iranian fig cultivars studied by Khadivi et al. (2018), fruit ostiole width ranged
from 0.00 to 10.8 mm. Also, in Iranian wild edible fig accessions studied by
Mirheidari et al. (2020), the range of ostiole width was 1.06–6.9 mm. In addition,
Darjazi (2011) reported that Fruits ostiol width was changed between 0.0 (Morabaii)
and 4.6 mm (Siah zoodras) in fig cultivars from Varamin province in Iran. Fruit
ostiole width has been reported as 2.34–8.79 mm with an average of 5.13 mm
(Aksoy et al., 1992), 1.50–4.00 mm (Koyuncu et al., 1998), 1.00–9.4 mm (Polat &
Ozkaya, 2005), 1.1–4.9 mm (Caliskan & Polat, 2008), and 2.25–8.93 mm (Gozlekci,
2011) in different fig growing areas in Turkey. Also, fruit ostiole width has been
reported as 0.00–8.20 mm (Hssaini et al., 2019), 0.60–9.10 mm (Hssaini et al.,
2020c), in different fig accessions from Morocco. In addition, Hssaini et al. (2020a),
who studied eleven fig varieties from Northern Morocco, reported that the ostiole
was generally large fruits such as ‘Kadota’ and ‘Palmeras’ (7.69 and 5.81 mm,
respectively).
Fruit Skin Cracks Cracking around the ostiole is undesirable because the patho-
gens and pests enter fruit via these cracks (Can, 1993). Skin cracks influence con-
sumer acceptance because consumers think that this character indicates the degree
of fruit ripening (Mahmoudi et al., 2018). Many factors (i.e., genetic, morphologi-
cal, environmental, and physiological) influence fruit cracking. Under the same
environmental conditions, fruits from different cultivars show differences in crack-
ing susceptibility. Some correlations have been observed between susceptibility of
fruit cracking and some fruit traits (fruit shape, fruit size, fruit firmness; anatomy
and strength of the fruit skin, stomata in fruit skin, cuticular properties, osmotic
concentration, water capacity of the fruit pulp and growth stage of the fruit). Also,
orchard management (such as irrigation and nutrition) and environmental condition
(such as temperature, wind, and light) can influence fruit cracking. Besides, fruit
cracking is a quantitative trait controlled by several genes. Therefore, the best way
to reduce fruit cracking would be suitable for orchard management that takes into
account and minimizes the stress of the water, nutrition, and physiological factors
that contribute to fruit cracking. Also, the most resistant cultivars to fruit cracking
that have desirable fruit quality can be selected for cultivation (Khadivi-Khub,
2015). Fruit splitting in figs is due to sudden changes in the internal pressure of the
fruit that can be the result of cool temperatures and high humidity while the fruit
matures, precipitation at fruit maturity, or over-caprification (excessive pollination)
(Crisosto et al., 2011).
In Iranian fig cultivars studied by Khadivi et al. (2018), fruit cracking percentage
varied from 0.00% to 70.00%. The seven genotypes, including ‘Siah-1’,
‘Siah-2’,’Sabz-22’, ‘Sabz-40’, ‘Sabz-60’, ‘Choopani’, and ‘Mati’ had no fruit
cracking. The highest width of the cracked fruit section was 9.10 mm and was
observed in the ‘Sabz-71’ genotype, while the lowest value of this trait was 0.00 mm.
Also, in 92 Iranian wild edible fig accessions studied by Mirheidari et al. (2020),
fruit skin cracks were not observed in the majority of accessions (72), and among
6 Phenotypic Variability of Fig (Ficus carica L.) 151

the accessions having fruit skin cracks, the cracking area was predominantly on the
base of fruits (16). Furthermore, Hssaini et al. (2019) reported that the skin cracks
were primarily absent in local fig clones from Northern Morocco.
Fruit Flesh and Skin Thickness and, Ease of Peeling The thickness of the flesh
and its hydration rate can effectively crack the skin. In Moroccan fig accessions
studied by Hssaini et al. (2020c), fruit peel thickness (mm) was 1.32–8.35. Hssaini
et al. (2020a), by evaluating 11 local and introduced fig cultivars cultivated in the
Moroccan climate, reported that ‘Breval Blanca’ and ‘Nabout’ cultivars had the
thick-skinned fruit, whichthe average values were, respectively, 8.35 and 5.91 mm.
Also, the Ease of peeling is critical to satisfying local and global customer pref-
erences (Caliskan & Polat, 2008; Can, 1993). Easy skin peeling is an essential char-
acteristic of consumer acceptance (especially for the cultivars that have thick skin,
such as ‘Bakkor Biadh’ and ‘Onk Elhamam’) because consumers tend to peel figs
before eating (Mahmoudi et al., 2018). Interesting cultivars in fig cultivars grown in
Central-Western Spain, with respect to fruit quality were ‘Antigua’, ‘Cuarterón’,
‘Cuello de Dama Blanco’, ‘Gota de Miel’ and ‘Pringo de Mel’. These developed
very juicy and easily peeled fruits (Pérez-Sánchez et al., 2016).
Fruit Skin and Flesh Color Fruit skin and flesh colors determine ripening time.
These characteristics are also used with other features to determine the promising
fig genotypes in breeding studies and the selection of accessions used in breeding
studies (Tsantili, 1990; Sacks & Shaw, 1994). In addition, the fruit skin color of
fresh figs is essential for consumer preferences (Mirheidari et al., 2020), and fresh
figs with pink and red flesh colors are preferred by consumers (Caliskan & Polat,
2008). The peels and pulps of different cultivars had variable levels of polyphenols,
flavonoids, anthocyanins, tannins, and antioxidant activity. Generally, fig fruit peels,
especially those with a dark color, contained the highest concentrations of phyto-
chemicals and exhibited the highest antioxidant activity compared to fig fruit pulps.
Based on mineral and phytochemicals, it is recommended consumption of the whole
fig fruit (Mahmoudi et al., 2018).
Fig skin color varies from dark purple to green. According to their color, figs are
categorized into 3 groups: White, Red, and Black. The skin of the white variety is
white, yellowish, or even green, e.g., the Kadota variety. Another variety known as
Dotato has yellowish-green skin and purple-colored flesh. The white variety is
either seeded or colored seedless. The reddish variety is distinguished through its
bluish brown color. The black variety includes those figs with dark red to black skin
color. This group includes varieties like Brown Turkey, Celeste, and Sari Cob
(Stover et al., 2007b). Gozlekci (2011) indicated that fruit skin color in fig acces-
sions ranged from green-yellow to black among fig accessions from Turkey. In 92
Iranian wild edible fig accessions studied by Mirheidari et al. (2020), fruit skin
ground color was strongly variable. It included light violet (3 accessions), violet
(20), dark violet (35), red (5), yellow (7), yellow-violet (7), yellow-red (1), violet-
cream (6), cream (6), and light brown (2) and internal pulp color included cream (1),
gold (1), red-gold (6), light red (3), red (7), dark red (13), light violet (8), violet (29),
152 A. Khadivi and F. Mirheidari

light brown (16), brown (6), and dark brown (2). Also, Darjazi (2011) reported that
the fruit skin color varied from yellow to black in fig cultivars from Varamin prov-
ince in Iran. ‘Paizeh’ and ‘Morabaii’ cultivars had canary yellow colors. ‘Zard’
cultivar was yellow-green, and ‘Bidaneh’ cultivar was light green, while ‘Hallavi
riz’ and ‘Hallavi dourosht’ cultivars were light-brown, ‘Siah bolbolriz’ cultivar was
purplish-brown, ‘Siah zoodras’ cultivar was purple and ‘Siah diras’ cultivar was
black. Also, in their study, the skin ribs were Dark-brown (for ‘Hallavi riz’ and
‘Hallavi dourosht’) and absent for the remaining cultivars.
In local fig clones collected in Northern Morocco by Hssaini et al. (2019), the
skin colors of sampled fruits varied from the blue-purple to the green-yellow, and
the predominant internal pulp color was dark-red and purple. Also, fruit flesh color
in fig cultivars grown in Central-Western Spain ranged from white to red (Pérez-
Sánchez et al., 2016).
In 82 Lebanese fig accessions studied by (Chalak et al., 2008), skin ground color
ranged from a purplish black to yellow with all the intermediate scales. The skin
over color was absent or present as irregular patches with yellow, purple, or green
sectors. Internal color varied between white, amber, pink, dark pink, red and dark
red. In Iranian fig cultivars studied by Khadivi et al. (2018), Fruit skin color was
most often yellow (40 genotypes, 46%) and followed by dark yellow (31 genotypes,
35.6%), and also the fruit pulp internal color was yellow in most of the genotypes
(63 genotypes, 72.4%). In 38 Slovenian fig accessions described by Podgornik et al.
(2010), the fruit skin ground color of the main crop varied from black for
‘Kamberij-1’ to purple for ‘Flazana-2’, ‘Crni Matalon’, ‘Crnica’, ‘Miljska figa’ and
‘Kamberij-2’ through purple-brown for ‘Madona Rjava’ and ‘Rjavi Matalon’,
brown for ‘Kanor’, dark green for ’Crna Petrovka’ and ‘Petrovaca crna’ to green
and yellow-green for all others varieties and the internal color varied from amber to
dark red. Eleven accessions were dark red, nine accessions were pink, and only
three accessions were amber-colored.
Fruitlet Amount, Size and Weight In Iranian fig cultivars studied by Khadivi et al.
(2018), the ‘Mati’ genotype was seedless, while the range of seed number in other
genotypes ranged from 24 (in ‘Siah-1’ and ‘Siah-2’ genotypes) to 575 (in ‘Sabz-11’
genotype).
Also, Darjazi (2011) reported that the number of seeds per fruit was high (for
‘Zard’), medium (for ‘Paizeh’, ‘Siah diras’, ‘Hallavi dourosht’), and low for the
remaining fig cultivars from Varamin province in Iran. The fruit of ‘Bidaneh’ had
the strongest aromatic flavor. In 92 Iranian wild edible fig accessions studied by
Mirheidari et al. (2020), the seed amount was high in most of the accessions (61).
The seed length, seed width, and seed thickness range were 1.21–2.13 mm,
1.00–1.90 mm, and 0.69–1.52 mm, respectively. Hssaini et al. (2019) reported that
most of the genotypes from Northern Morocco had a medium seed size.
6 Phenotypic Variability of Fig (Ficus carica L.) 153

5.3.1 Environmental Condition’s Effects on Fruit Characteristics

Climate markedly affects figs’ size, shape, skin, and pulp color (Condit, 1947).
Cooler climates produce greener than yellow skins, more vivid pulp colors, and
larger, more elongated fruits. Crane (1986) has suggested that the larger individual
size of the first breba crop, which competes with shoot growth and the second crop
for available carbohydrates, is due to its development during a cooler period. In
addition, climate may also affect pollination requirements. Other environmental
conditions such as rain, hail, and wind can reduce fruit quality and production. Rain
may cause fruit splits. Splitting results from sudden changes in the internal fruit
pressure brought on by cool temperatures and/or high humidity as the fruit matures
(Ferguson et al., 1990). Splitting in ‘Calimyrna’ and other varieties may also result
from excessive pollination and the growth of too many developing seeds during fruit
ripening. Strong winds during the ripening season whip the foliage and cause scar-
ring of fruits such as ‘Kadoti’ and ‘BrownTurkey’ (Flaishman et al., 2008a).
A comparison of the role of climatic conditions in dry and fresh fig production
revealed that dry fig production is strongly dependent on climatic conditions and is
successful mostly under dry and warm-temperate climates. Fresh figs, however, can
be cultivated under a wider range of ecological conditions (Sahin, 1998).
Spontaneous fig trees can potentially exchange genetic material each year
(Kjellberg et al., 1987). A study of enzymatic polymorphism revealed that the spon-
taneous fig trees of southern France show that alleles are not randomly distributed;
their frequencies vary regularly with the climate and reflect the role of adaptation
(Kjellberg & Valdeyron, 1984).
The fig weight (20–60 g) in a study quoted by Crisosto et al. (2011) on ten fig
varieties (152-4s, ‘Brown Turkey’, ‘Kadota’, ‘Mission’, ‘Orphan’, ‘Panachee’,
UCR 276-14, UCR 291, ‘White Texas Everbearing’ and ‘Zidi’) grown under
California conditions, was comparable with that of Turkish varieties [22–52 g
(Caliskan & Polat, 2008), 40–65 g (Bostan et al., 1998), and 30–90 g (Ozeker &
Isfandiyaroglu, 1998)]. The varieties grown in California had similar fruit length
(41-63.5 mm) to varieties from Turkey (38.5–62.0 mm) but were smaller in fruit
diameter (35.6–48.4 mm versus 45.0–55.0 mm) (Bostan et al., 1998). Soluble solids
concentrations of these varieties grown in California (13–31%), though with more
extreme values, were in general comparable to the Turkish varieties [15.1–21%
(Bostan et al., 1998), 16–27% (Ozeker & Isfandiyaroglu, 1998), and 20.1–27.4%
(Caliskan & Polat, 2008)]. Finally, the TA of the varieties grown in California
(0.14–1% citric acid) was, in general, higher than the same varieties from Turkey
[0.09–0.26% (Caliskan & Polat, 2008), 0.14–0.22% (Bostan et al., 1998), and
0.06–0.15% (Ozeker & Isfandiyaroglu, 1998)].
154 A. Khadivi and F. Mirheidari

5.3.2 Caprification Effects on Fruit Morphological Characteristics

Caprification is the pollination of long-styled female flowers of the edible fig by

wasps carrying the pollen from the profichi borne on the caprifig (hermaphroditic,
with male and female flowers in the profichi fruits). The number of male figs dis-
pensed and length of pollination period depends on weather conditions (Condit,
1947), age, size, the yield of female cultivar, and type of caprifig.
The common practice of caprification is to distribute the male figs at intervals of
a few days over about 3 weeks (Khadari et al., 1995b; Rahemi & Jafari, 2008; Zare,
2008). In Tunisia, two to six male figs are connected with a wire, or a stick passed
through their neck and hung onto branches of female fig trees of the Smyrna-type
(Mars et al., 2008). In California, ripe caprifigs are cut and placed in bags or baskets
that are stapled on female trees of the Smyrna and San Pedro-type (Stover et al.,
2007a). In Iran, caprifigs are put inside containers hung on the female fig trees of the
Smyrna-type (Zare, 2008).
The selection of pollen sources with good quality and quantity of pollen is an
important and essential practice in fig orchards (Küden & Tanriver, 1998; Khadivi-
Khub & Anjam, 2016). In addition, it is well known that the type of caprifig can
have a significant effect on the skin color, ostiole width, total soluble solids, time of
ripening, seed germination, size and shape of syconia (Janick & Moore, 1975;
Rahemi & Jafari, 2008; Gaaliche et al., 2011).
1. Caprification and Fruit-Set and Yield Fig caprification is a complex system for
fruit production and is a model system for studying the mutualism and coevolution
between figs and pollinating wasps (Bandelj et al., 2007; Giraldo et al., 2008a).
Despite its importance, this mechanism remains unclear, and the classification of fig
types based on pollination and fruit production is still ambiguous. Ferrara et al.
(2016) reported that varieties such as Rosso Comune, Nero Terlizzi, Nero Sava, etc.,
clearly needed profichi (wasps) to set figs and obtain good yields. Other varieties in
their study, such as Dottato Marchese, Dottato Sava, Nero Crotone, Regina Gioia,
Rosso Triggiano, etc., were not mainly influenced by showing satisfactory fruit-set
and yield with or without caprification. Their results indicated that fruit-set was
significantly influenced by the presence/absence of profichi in the orchard (i.e., cap-
rifigs), with varieties producing well yield only when caprificated and varieties set-
ting figs parthenocarpically, thus not strictly needing profichi for the pollination.
The fruit set of brebas was highly variable, with some varieties setting in significant
numbers of fruits (Dottato Marchese, Dottato Sava, Monaca, etc.) and others show-
ing potential for brebas production Petrelli, Sant’Antonio, Rosso Triggiano, etc.
(Ferrara et al., 2016).
Pollination by caprification greatly influences fruit size, which can double, while
the fruit set is also noticeably increased (Gaaliche et al., 2011). Ferrara et al. (2016)
indicated that yield was dramatically reduced when figs were non-caprificated
(either manually or naturally). Indeed, non-caprificated figs show an internal cavity,
dry matter content reduction, a collapse of pedicel development, and lack of seeds,
and the fruit is significantly firmer at harvest (Trad et al., 2013a). Physiologically,
6 Phenotypic Variability of Fig (Ficus carica L.) 155

caprification acts indirectly as a precursor of several hormone activities.

Pourghayoumi et al. (2012) showed a significant difference in the time of fruit rip-
ening in Iranian cultivars, but the effects of the pollen sources on the time of fruit
ripening were not significant. Their results were in contrast to the results found by
Rahemi and Jafari (2008). It is well known that ethylene plays a significant role in
ripening (Galil, 1968). Fertilized seeds produce a large amount of auxin, which may
stimulate ethylene production within the tissue (Ryugo, 1988). Ethylene, responsi-
ble for physiological and biochemical changes in fruit development and maturation,
may be enhanced by pollination. Ferrara et al. (2016) data on caprificated and non-
caprificated fruits are quite similar to the percentages reported by others (Rahemi &
Jafari, 2008; Zare, 2008). This demonstrated that caprification is necessary for San
Pedro and Smyrna type and Common type varieties. Whether required or not, cap-
rification can increase fruit-set and improve several organoleptic quality aspects
(Trad et al., 2013a). Even though caprification enhances fig production and yield,
little information is available on the genetic basis of this complex and exciting sys-
tem. Large-scale comparative gene expression studies have provided comprehen-
sive data on its effect on fruit physiology and elucidated genetic factors underlying
the traits involved in type differentiation (Ikegami et al., 2013).
2. Caprification and Fruit Morphometric Characteristics Pollinated figs are usu-
ally considered larger greener, and darker interior pulp color than unpollinated figs
(Condit, 1947; Oukabli et al., 2003; Michailides et al., 2008). Flaishman et al.
(2008a) reported that the caprified fruits of cultivar ‘Autumn Honey’ were bigger,
with darker purple skin color and red pulp, and have longer storability than non-
caprified fruits. In Tunisian cultivars, fruit weight, length and diameter, ostiole
diameter, flesh thickness, and the number of fertile seeds were significantly increased
by pollination rate (Gaaliche et al., 2011).
Pourghayoumi et al. (2012) confirmed that fruit length was significantly affected by
pollen sources and female cultivars, and also, fruit diameter was significantly
affected by female cultivars. Their results are in parallel with those reported by
Rahemi and Jafari (2008) and Gaaliche et al. (2011). In addition, these findings
agree with those of Bose and Mitra (1990), who indicated that the length of the fruit
neck of Smyrna-type figs was affected by pollen sources. The specific pollens with
increased IAA (indole-3-acetic acid) may affect the cell number in early fruit devel-
opment and stimulate cell elongation (El-Hamady et al., 2010; Khan &
Chaudhry, 2010).
Pourghayoumi et al. (2012) found that pollen sources did not significantly affect
fruit weight and ostiole width. Due to fruit weight, their results follow other reports
as in Iran; pollen sources had no significant effect on fruit weight (Rahemi & Jafari,
2008). However, the results contrast the results found by Gaaliche et al. (2011).
Pollen source affects the growth of ovarian tissues by stimulating hormone produc-
tion and exerting a specific effect on fruit growth (Shafique et al., 2011). Due to
ostiol width, the results of other reports in Iran and Tunisia confirmed that pollen
source had a significant effect on ostiol width (Rahemi & Jafari, 2008; Gaaliche
et al., 2011).
156 A. Khadivi and F. Mirheidari

In other experimental works, caprification was evident in fruit size, weight, and
flesh thickness (Trad et al., 2013a). Caprified figs have almost twice the weight of
non-caprified fruits. Caprification increased fig weight with a proportional increase
in the fruit’s flesh. Increased flesh thickness undoubtedly improves the consistency,
taste, and flavor of figs when ripe Non-caprified figs showed a larger internal cavity
and lack of seeds and the fruit was significantly firmer at harvest time than caprified
fruits (Trad et al., 2013a).
3. Caprification and Dry Matter Content Pollination increases the dry matter con-
tent of the fruit (Trad et al., 2013a, 2014). This is an important fruit qualitative
parameter when figs are destined for drying (Piga et al., 2003). Pollen source affects
ovarian tissue growth and fruits probably by stimulating hormone synthesis. The
increase in the synthesis of indole-3-acetic acid may stimulate cell elongation
(Pourghayoumi et al., 2012). Thus, caprification may indirectly act as a precursor of
several hormonal activities (Trad et al., 2013a).

4. Caprification and Skin/Pulp Color It is well known that the type of caprifig can
significantly affect the color of both the fruit skin and its interior edible flesh (Janick
& Moore, 1975). Skin color and external visual appearance exhibited no change in
an experiment with deficient caprification. However, good pollination generally
leads to pronounced pulp color, and caprified figs are richer in anthocyanins (Trad
et al., 2013a). This suggests that biosynthesis of phenolic compounds occurs later
after pollination and can be affected by caprification (Trad et al., 2013a). Janick and
Moore (1975) reported that the type of caprifig might significantly affect the color
of both the fruit skin and edible flesh and the size and shape of syconia. In addition,
Bose and Mitra (1990), Rahemi and Jafari (2008), and Pourghayoumi et al. (2012)
indicated that pollen sources had a significant effect on skin color of fig fruit.

5. Caprification and Fruit Firmness Both caprified and non-caprified figs lose
their firmness, but caprification enhances fruit softening (Trad et al., 2013a).
Although some authors reported that total fibers were somewhat fewer in caprified
figs than in non-caprified fruits, Trad et al. (2013a) found that caprification did not
affect the dietary fiber content of figs. Caprified fruits, having better pollination,
contain less aborted flowers and more seeds. Thus, the development of the flesh
appeared to be proportional to the number of seeds (Trad et al., 2013a). Pectin, esti-
mated by the uronic acid concentration, was less abundant and had a higher meth-
ylation degree after pollination, while the fruit had softer textures (Trad et al., 2014).
Pollination may allow the proper ripening of fruits. Enzymes like polygalacturonase
or pectin-methyl esterase efficiently proceed to fruit softening (Gross & Sams,
1984). It is essential to highlight that caprification improves the fig receptacle’s
textural properties by increasing the pectin’s methylation degree. Fertile seeds
resulting from pollination may be involved in hormonal activity stimulating many
related enzymes of the wall matrix depolymerization, in particular polygalacturo-
nase (PG) and pectin methylesterase (PME) (Trad et al., 2014).
6 Phenotypic Variability of Fig (Ficus carica L.) 157

6. Caprification and Fruit Cracking and Splitting Excessive pollination increases

fruit splitting and incidence of internal defects incited by microorganisms (Stover
et al., 2007a). The high number of fertilized seeds may be a reason for ostiole crack-
ing (Crisosto et al., 2010). Zare (2008) showed that using ‘Daneh-Sefid’ as caprifig
to pollinate Iranian cultivar ‘Sabz’ increased the number and diameter of seeds,
resulting in a rapid ostiole-end cracking before ripening of the fruits. As a result, the
female trees closest to the caprifigs can be overfertilized, and their figs may split
(Flaishman et al., 2008a). In addition, Increase in fruit size and tissue softening
occurs during the ripening of fig fruit. Increased risk of splitting could be linked to
the cell wall compositions and/or to the relative ratio of peel, the resistant tissue, to
a pulp (Trad et al., 2013a). Softening is due to cell wall modification in both the
pectic and hemicellulosic polymers in the receptacle and drupelets tissues.
Quantitative changes in polysaccharides occurred in both tissues. Qualitative varia-
tions between the two tissues concerned only the pectic but not the hemicellulosic
polymers (Owino et al., 2004a, b). Fruit ripening is a complex metabolic event and
includes increased cell wall hydration, disassembly of cell wall polysaccharides,
reduction in cell-to-cell adhesion, and decline in turgor pressure. Eleven genes
encoding cell wall modifying enzymes during the ripening of the fig fruit (endo-
polygalacturonases (PGs), endo-glucanases (EGases), ß-galactosidases (ß-Gals),
xyloglucan endotransglycosylases (XTHs), expansins (Exps) and
α-arabinofuranosidases (Arabfs)) were reported (Owino et al., 2004b). These genes
may act together in an interdependent way to achieve softening. The average nitrate
content of persistent figs was much higher than that of non-persistent ones. The
regulation of indole-acetic acid oxidase is involved in the process, and persistent
cultivars are expected to have higher auxin levels (Flaishman et al., 2008b).

6 Descriptor’s Optimization Tools

Some optimization tools for descriptors that are empirically useful for selecting
more viable and reliable morphological variables and detecting the distinction
among the genotypes are analysis of correlation coefficients and principal compo-
nent analysis (PCA). In addition, phenotypic and genetic studies from a heritability
point of view can also help select more stable morphological characteristics in the
description and discrimination of genotypes. The higher the percentage of heritabil-
ity in a variable, the more stable and reliable the traits. However, traits with a low
percentage of genetic inheritance and those most affected by environmental condi-
tions can also be used to identify different ecotypes of a genotype/cultivar.
Morphological characterization is a complex process, and, to optimize it, descrip-
tor lists for a high number of species have been developed. For example, in fig, a
group of international experts developed and published a list of descriptors
(International Plant Genetic Resources Institute and Centre International de Hautes
Etudes Agronomiques Mediterraneennes, 2003) to facilitate the characterization
process of this species. However, the analyses of morphological traits in fig are even
158 A. Khadivi and F. Mirheidari

more complex than in most fruit tree species as a result of some particular charac-
teristics of this species, such as the frequent production of two different crops (breba
and main crop), the presence of four different types of leaves, or its complex floral
biology. Consequently, the current descriptors list available has 192 traits, of which
126 are qualitative and 66 are quantitative. This high number of traits and the need
to replicate observations for at least 2 years make phenotypic characterization in
this species a slow time-consuming, and resource-consuming process, especially in
developing countries with scarce economic resources (Giraldo et al., 2010).

6.1 Principal Component Analysis (PCA)

Principal component analysis (PCA) is a technique for reducing the dimensionality

of such datasets, increasing interpretability while minimizing information loss. It
does so by creating new uncorrelated variables that successively maximize vari-
ance. In other words, it is the process of computing the principal components and
using them to perform a change of basis on the data, sometimes using only the first
few principal components and ignoring the rest.
Principal component analysis (PCA) is used to identify patterns of variability
among the genotypes and the traits with the highest variation between cultivars and
the greatest impact on their separation in the data set (Iezzoni & Pritts, 1991).
The approach used in Giraldo et al. (2010) work has suggested selecting a subset
of highly discriminant qualitative traits by finding a correlation between variables
and cultivars based on qualitative trait frequencies and multivariate analysis fol-
lowed by PCA in fig that, in order of importance, are the following: (1) production
type: common type (bifera and unifera type), Smyrna, San Pedro, and caprifig type;
(2) morphology of the main crop and the breba crop [fruit shape, lenticels color and
abundance, skin ground color, overcolor (regular bands and irregular patches), pulp
and pulp cavity color, stalk color, skin firmness, and presence of male flowers]; (3)
morphology of five-lobed and three-lobed leaves: central lobe shape, petiole sinus
shape, petiole color; (4) morphology of entire leaves: leave shape, petiole sinus
shape; and (5) tree morphology: tree growth habit, size of the tree, degree of branch-
ing, and bark tubers. Most of the variables are related to fruit characteristics (breba
and main crops), highlighting the importance of correct fruit characterization to
identify fig accessions correctly.
Also, in another study by Mirheidari et al. (2020) on 92 Iranian accessions of
wild edible figs with a total of 55 quantitative and qualitative morphological and
pomological traits were used for phenotypic evaluations. The PCA showed that the
traits measured were subdivided into 15 independent components, explaining 78.9%
of the total variance. The PC1 was positively correlated with 13 traits, including leaf
length, leaf width, central lobe length, upper lateral vein length, hair density on leaf
upper surface, petiole length, petiole thickness, ripening time, fruit length, fruit
width, fresh fruit weight, fruit stalk width, and fruit flesh thickness that explained
21.3% of the total variance. The vegetative-related traits, including branching,
6 Phenotypic Variability of Fig (Ficus carica L.) 159

branch density, canopy density, and leaf density, loaded significantly on PC2 and
accounted for 7.29% of the total variance. The PC3 was correlated with the number
of main leaf venation, lobe number, upper lateral lobe depth, and upper lateral lobe
base width, which accounted for 6.40% of the total variance. These findings, in
most cases, corresponded with the previous edible fig morphological studies (Mars
et al., 1998; Saddoud et al., 2011; Darjazi, 2011; Gaaliche et al., 2011; Ciarmiello
et al., 2015; Khadivi et al., 2018; Hssaini et al., 2020b) that have reported the impor-
tance of morphological characterization as the main factor in discriminating and
assessing breeding materials of fig trees. Khadivi et al. (2018) have reported that the
pomological characteristics are essential to evaluate the variation in traits of edible
fig accessions.
Similarly, many studies revealed that principal component analysis in qualitative
and quantitative morphological traits (vegetative and pomological) is very helpful in
evaluating the phenotypic diversity and correctly identifying fig accessions.

6.2 Correlation Coefficients

Overall, the inventory of plant material based on morphological traits is important

for managing genetic resources, maintaining the existing genetic variability, and
establishing a germplasm collection (Podgornik et al., 2010). Authors stress the
relevance of morphological assessment in the selection and breeding programs of
cultivars and claim such analysis should be performed before biochemical or molec-
ular studies are carried out (Djordjević et al., 2014; Khadivi et al., 2018; Khadivi-
Khub et al., 2012; Podgornik et al., 2010).
The correlation coefficient is a statistical measure of the strength of the relation-
ship between the relative movements of two variables. In other words, it reflects
how similar the measurements of two or more variables are across a dataset. The
values range between −1.0 and 1.0. A calculated number greater than 1.0 or less
than −1.0 means an error in the correlation measurement.
The correlation coefficients can provide information on the most important traits
in assessing genotypes (Norman et al., 2011; Khadivi-Khub & Anjam, 2016). In
addition, the correlation coefficients may provide information on the descriptors
that are potentially important in assessing fig genotypes (Norman et al., 2011).
Correlated variables can also be used to predict other ones and could be considered
of importance for the characterization and the discrimination and fig genotypes
(Podgornik et al., 2010; Khadivi-Khub & Anjam, 2016; Hssaini et al., 2020a). In
addition, a close relationship between traits could facilitate or hinder gene introgres-
sion since strong selection for a desirable trait could favor the presence of another
desirable trait from germplasm (Dicenta & Garcia, 1992).
The results reported in previous female fig studies have suggested that fruit and
leaf traits are important factors in differentiating and analyzing breeding materials
dealing with the morphological characterization of fig (Mars et al., 1998; Hedfi
et al., 2003; Saddoud et al., 2008b, 2011; Gaaliche et al., 2012; Aljane et al., 2012;
160 A. Khadivi and F. Mirheidari

Caliskan & Polat, 2012; Khadivi-Khub & Anjam, 2014; Khadivi et al., 2018;
Hssaini et al., 2020b; Mirheidari et al., 2020). Also, Papadopoulou et al. (2002) and
Almajali et al. (2012) reported that the fruit size and shape, ostiole width, leaf char-
acteristics, and leaf shape are very important for genotype selection. Therefore, fac-
tor analysis had great potential to differentiate the highlighted distinctions between
studied genotypes and agreed with the findings of others in the study of fig (Mars
et al., 1998; Saddoud et al., 2008b, 2011; Gaaliche et al., 2012). Correlations
between characteristics revealed by the PCA method may correspond to a genetic
linkage between loci of controlling traits or a pleiotropic effect (Iezzoni &
Pritts, 1991).
Among the examined morphological and pomological characters of 87 edible fig
genotypes in Iran by Khadivi et al. (2018), leaf length showed positive correlations
with leaf width, lobe number, central lobe depth, lateral lobe depth, petiole length,
and petiole thickness, and agreed with the finding of Khadivi-Khub and Anjam
(2014) in caprifig. The existence of close positive correlations among leaf traits
indicates that more leaf expansion leads to stronger aerial growth (Khadivi-Khub &
Anjam, 2014, 2016; Anjam et al., 2017). Also, in their study, fruit yield showed
positive correlations with leaf density, dried fruit width, and dried fruit weight. In
addition, dried fruit weight was positively correlated with leaf density, leaf length,
leaf width, leaf color, vein number, central lobe depth, and lateral lobe depth, and
also highly correlated with dried fruit length, and dried fruit width, following the
results of Khadivi-Khub and Anjam (2014, 2016) in caprifig. Khadivi et al. (2018)
found that fruit cracking percentage showed negative correlations with leaf color,
fruit fall type, and fruit skin color, while it showed positive collations with fruit
yield, ostiole width, and fruit cracking width, seed number, and seed weight. Also,
fruit cracking width was negatively correlated with fruit fall type, while this trait
indicated positive correlations with ostiole width, seed number, and seed weight.
Also, fruit quality was positively correlated with leaf density, dried fruit width,
dried fruit weight, ostiole width, fruit cracking width, and fruit cracking percentage
(Khadivi et al., 2018).
Among the examined characters for fruits of 22 cultivars in South Tunisia by
Mars et al. (1998), fruit weight was highly correlated with fruit diameter and flesh
thickness. Fruit diameter was also correlated with stalk diameter and flesh thick-
ness. Neck length was correlated to fruit length. Ostiole type and ostiole diameter
were also well correlated. Also, in Iranian fig cultivars studied by Khadivi et al.
(2018), dried fruit weight was positively correlated with leaf density, leaf dimen-
sions, dried fruit length, and dried fruit width.
According to five-year data of Turkish cultivars, fruit weight was found to have
positive correlation by fruit diameter (r = 0.92; P < 0.01), fruit length (r = 0.81;
P < 0.01), neck length (r = 0.35; P < 0.01), ostiolium width (r = 0.23; P < 0.01),
trunk diameter (r = 0.26; P < 0.01), fruit skin cracks (r = 0.26; P < 0.01) and nega-
tive correlation by TSS (r = −0.26; P < 0.01) and fruit ribs (r = −0.21; P < 0.01). It
was also found out in that period that as the trunk diameter increased, so did the fruit
size while the ostiolium width was smaller. This could be related to the increase in
the fruit size with age and the bigger trunk diameter (Polat & Caliskan, 2017).
6 Phenotypic Variability of Fig (Ficus carica L.) 161

Indeed, Botti et al. (2003) reported that fruit size could continue to grow until the
tree gains its final shape.
Hssaini et al. (2020b) confirmed that Fruit weight was positively correlated to
stalk width, neck length, width, ostiole width, and peel thickness. These results
were similar to those reported in several works (Mahdavian et al., 2006; Gozlekci,
2011; Darjazi, 2011; Gaaliche et al. 2011; Aljane et al., 2012; Polat & Caliskan,
2017. Khadivi et al., 2018). In addition, they concluded that all fruit dimensions
were positively and strongly correlated to fig volume except for stalk length and
ostiole width.
Hssaini et al. (2019) concluded that total soluble solids (TSS) are negatively and
significantly correlated to fruit weight, width, and the volume, which means that
cultivars with high content of total soluble solids would tend to present a reduction
in the fruit weight, width, and volume. Similar results were demonstrated in figs
(Gozlekci, 2011) and other fruits such as yellow passion (Viana et al., 2003).
In the past decades, strong attention has been given to combining several descrip-
tors of interest as an important quality parameter in fig selection. Morphological and
biochemical traits seem to have attracted particular substantial interest since some
of them can be used to predict each other based on correlations. In addition, it can
provide information about the relationships between variables that are potentially
important in assessing fig genotypes (Hssaini et al., 2020a).

6.3 Heritability

The main role of heritability is related to the fact that it expresses the reliability of
the phenotypic value as an estimator of the genotypic value, such that the higher the
heritability in selection (Acquaah, 2012). Fruit weight and size (diameter and
length), SSC, TA, and SSC:TA ratio, skin color (luminosity, chroma, and hue), eth-
ylene production and respiration rates (CO2 production), firmness, ostiole diameter,
internal cavity diameter, shriveling, growth cracks, shelf life and susceptibility to
blemishes are affected by genotype (Crisosto et al., 2011).
Although the findings of Cowart and Graham (1999) indicated that the length
and width of the leaves are not good biometric characteristics, it is worth noting that
higher values of these traits in combination with other characteristics can help deter-
mine possible selection characters. For example, Castellen et al. (2007), character-
izing 68 accessions of the Active Germplasm Bank of Papaya of Embrapa Mandioca
and Tropical Fruit through multivariate analysis, detected that the components 1
(petiole color) and 2 (leaf length) explained most of the total observed genetic varia-
tion (67.80%). Also, in fig accession studied by Rodrigues et al. (2019b), despite the
low heritability observed for length and width of the leaves, the number of leaf lobes
showed high heritability (0.85), with a CVg/Cve ratio higher than 1. The values
observed for the number of leaf lobes characters ranged from three non-pronounced
to seven well-defined lobes; these extremes were observed in ‘Troyano’ and
‘Brunswich’ accessions, respectively.
162 A. Khadivi and F. Mirheidari

7 Conclusions

Traditionally, the amount and distribution of genetic diversity have been determined
by morphological markers. Although environmental conditions and agronomic
technologies strongly influence the expression of these markers, the morphological
characterization is a highly recommended first step that should be made before
more in-depth biochemical or molecular studies are attempted (Hoogendijk &
Williams, 2001).
The fig tree has not been subjected to intensive plant breeding programs, and
thus many fig tree populations exhibit rich genetic biodiversity that can only be fully
exploited once it is properly identified and classified. Traditionally, the plant germ-
plasm characterization with the aim of its conservation has been carried out using
morphological or agronomical traits. Nevertheless, despite the progress in elaborat-
ing descriptors, fluctuations among years, environments, or repetitions have made
application difficult until recently (Giraldo et al., 2010). However, selecting highly
discriminant variables is important to optimize resources for a feasible morphologi-
cal characterization. This is especially important in a crop such as a fig fruit with
hundreds of genotypes described worldwide in which many synonymies and hom-
onymies may be observed (Khadivi et al., 2018).
Presumably, the most reliable method for accurate investigation and optimizing
resources of reliable and highly discriminant variables for a feasible morphological
characterization and to detect separation of genotypes (varieties, cultivars, and
accessions) is vegetative propagation and thus cloning of a homogeneous and uni-
form genotype and planting them in different regions with different climatic condi-
tions and of course with a specific nutritional program. Then, under different
environmental and geographical conditions, it is possible to diagnose stability and
instability in the measured qualitative and quantitative morphological traits in that
cultivar. In this way, the altered and environmentally affected traits can be ignored,
and as a result, valid and stable morphological and distinctive traits in that cultivar
can be obtained. Also, in this method, we can define a pattern for unstable traits in
different climatic conditions as additional side information for different cultivars.

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Chapter 7
Morpho-Chemical Characteristics Useful
in the Identification of Fig (Ficus carica L.)
Germplasm

Oguzhan Caliskan, Safder Bayazit, and Derya Kilic

Abbreviations

CIHEAM International center for advanced Mediterranean agronomic studies

IPGRI International plant genetic resources institute
PCA Principal component analyzes
TSS Total soluble solids
UPOV The international union for the protection of new varieties of plants

1 Introduction

Fig is one fruit species with a history as much as human history. The spread of the
figs from Anatolia, the Caucasus, where it originated, to the whole world is accepted
as a sacred fruit in the monotheistic religions, a fruit that is fondly consumed, and
its nutritional properties. However, after this spread, important morphological dif-
ferences emerged both in the origin areas and in the areas to which it was moved
(Çalışkan & Dalkılıç, 2022; Condit, 1947). Figs can be grown in tropical and sub-
tropical regions, although it is cultivated mainly in the mild temperate climate of the
Mediterranean basin. The fig is probably the only edible fruit in a family of many
giant rubber tree species growing in tropical and subtropical climates and has, there-
fore, been traded for thousands of years. It is considered an exotic fruit, especially
in countries where cultivation is not possible (Aksoy et al., 2007).
For ages, the fig tree’s fruit has been valued in its fresh and dry forms. Seventy
percent of the world’s fig production is produced in the countries along the
Mediterranean coast. In these countries, figs are an essential component of the

O. Caliskan (*) · S. Bayazit · D. Kilic

Department of Horticulture, Faculty of Agriculture, Hatay Mustafa Kemal University,
Hatay, Turkey
e-mail: [email protected]; [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 175
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_7
176 O. Caliskan et al.

Mediterranean diet, which is considered one of the healthiest in the world and asso-
ciated with long life (Caliskan, 2015; Solomon et al., 2006). Especially the polyphe-
nol and antioxidant contents of black-colored figs are richer than the light-colored
ones, increasing the interest in colored individuals because they are healthier
(Caliskan & Polat, 2011).
Ficus species are gynodioecious, bearing either hermaphroditic or edible figs on
separate plants. Figs have different pollination biology from other fruit species.
Male fig plants, caprifigs, produce male and short-styled (gall) female flowers in
syconium (Fig. 7.1). Edible figs produce only female flowers (Beck & Lord, 1988),
but their pollination requirement differs. The female flowers in edible figs are long-
styled and occur as succulent fruitlet (Stover et al., 2007).
Three types of female figs are defined based on their cropping. The common fig
develops fruit parthenocarpically without pollination and could produce one (unifera
cultivars) or two (bifera cultivars) crops. The Smyrna-type fig requires pollination
with pollen from caprifigs. Furthermore, the San Pedro-type fig produces a first crop
parthenocarpically (breba) and a second main crop of fruit only after pollination
with pollen from caprifigs (Flaishman et al., 2008). Condit (1955) indicated that
78% are common types, less than 4% are San Pedro types, and the remaining 18%
are Smyrna types of edible figs. Besides, cultivars are diverse in such properties as
leaf morphology, plant vigor, fruit skin and internal color, fruit flavor, total soluble

Fig. 7.1 The appearance of ripe fruits and fruit flesh differences in caprifig (left) and edible
fig (right)
7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 177

solids content, titratable acidity, fruit shape, skin thickness, ostiole width, and dura-
tion of fruit production (Flaishman et al., 2008).
A caprifig tree gives three crop periods of fruit each year. Profichi fruit in sum-
mer, mammoni fruit in fall, and mamme fruit in winter. The profichi, the main cap-
rifig crop, comes across with the main summer crop of edible fig trees. The caprifig
trees supply a pollen source for caprification, which is the transfer of pollen from
caprifig trees to edible fig trees via the wasp vector Blastophaga psenes. Caprification
is a traditional practice in all fig-growing regions and is essential in edible figs’ fruit
set and quality (Condit, 1947; Ferrara et al., 2016; Caliskan et al., 2017a).
The fig fruits, primarily used for fresh consumption, have various colors, sizes,
shapes, and flavors in origin areas. The names of these figs are mainly given based
on local geographic origin, fruit size, fruit shape, fruit skin color, ostiole color,
maturity dates, or the name of the orchard owner. Researchers may encounter simi-
lar synonym and homonym genotypes because of the interchange of fig plant mate-
rial from nearby locations (Caliskan & Polat, 2012a).
The fig plants remain mostly as traditional crops as in other Mediterranean coun-
tries, and important genetic variations have been taking place due to biotic and
abiotic processes such as urbanization, the extension of intensive crops, and fig
mosaic disease (Salhi-Hannachi et al., 2004). Losses in the genetic diversity of crop
species due to commercialization have led to the need to preserve the present genetic
resources as much as possible, not only for the long-term survival of the species but
also to ensure enough variability for breeding programs (Esquinas Alcazar, 2005).
There has been some research dealing with the genetic diversity in edible fig germ-
plasm (Aksoy et al., 2003; Giraldo et al., 2010; Podgornik et al., 2010; Şimşek &
Yildirim, 2010; Stover & Aradhya, 2008); however, to date, there has been little
research into the genetic resources available for caprifig germplasm (Dalkılıç et al.,
2011; Khadivi-Khub & Anjam 2014; Caliskan et al., 2017b).
Plant genetic resources should be identified and protected due to their advan-
tages, such as being compatible with different ecological environments and contain-
ing genes resistant to different diseases and pests. For this reason, in addition to the
determination of morphological, phenological, and pomological data of genotypes,
the development of technology, the popularity of gene science, and the use of
genetic studies, especially DNA markers, to identify individuals have led many
developed countries to carry out genetic studies to identify their plant gene resources.
Characterization designs of morphological variability within the collections and
selection of the most significant variables should be carefully preserved to conserve
better and utilize genetic resources (Giraldo et al., 2010). This characterization is a
highly recommended first step before starting biochemical or molecular studies
(Hoogendijk & Williams, 2001). Recent developments show that the most impor-
tant natural resource of the current century plants genetic resources. Protecting
these resources necessitates preserving the genetic materials as they are today and
transforming these resources into benefits. Improving the use of plant genetic
resources for food and agriculture can be achieved by determining the essential
properties of the material during preservation (Inal, 2002).
178 O. Caliskan et al.

In recent years, principal component analysis (PCA) has been used to identify
fruit genetic resources and in different stages of breeding programs. This method
determines the most effective ones among hundreds of plant characteristics and
saves time and labor by reducing the number of examined attributes (Giraldo et al.,
2010; Saddoud et al., 2008; Simsek et al., 2020). This chapter presents the results of
the studies examining the morpho-chemical properties of figs using the principal
component analysis.

2 Plant and leaf Characteristics

The first detailed study on the plant and leaf description of fig genetic resources was
carried out by Condit (1941). He identified more than 600 fig genotypes, especially
leaf and fruit characteristics (Condit, 1955). In the following years, the characteris-
tics were evaluated by different researchers, and a fig descriptor was formed by
IPGRI and CIHEAM (2003). In addition, another fig descriptor was published by
UPOV in 2007. In both fig descriptors, nearly 200 features, including all plant and
fruit quality characteristics used to identify fig genetic resources, have been defined,
and expensive processes make the characterization of large germplasm collections
difficult. Therefore, useful variables in distinguishing fig genotypes have been
determined (Table 7.1).
The majority of the edible figs have an open and spreading canopy for tree growth
(Condit, 1941; Giraldo et al., 2010). Furthermore, the leaf shape is mainly G with
three lobes or C with five lobes (Caliskan & Polat, 2012a). The plant and leaf char-
acteristics that effectively distinguish the genotypes of edible figs are presented
(Table 7.2). The number and shape of lobes per leaf (Saddoud et al., 2008), tree
growth habit, size of the tree, degree of branching (Giraldo et al., 2010; Pérez-
Sánchez et al., 2016), terminal bud length and diameter (Essid et al., 2017), leaf
length, leaf width, leaf area, the density of hairs/spicules on the leaf’s upper surface,
and petiole thickness (Caliskan & Polat, 2012a; Ciarmiello et al., 2015; Khan et al.,
2022; Kokaj, 2021; Mirheidari et al., 2020; Podgornik et al., 2010) are the traits
used for the discrimination of edible figs. Although there are different leaf forms on
the same shoot, leaf shape is an essential morphological criterion in identifying fig
genotypes. In addition, apical dominancy, the number of leaves per shoot, the num-
ber of fruit per shoot (Caliskan et al. 2012a; Kokaj, 2021), and the beginning of fruit
maturation (Mirheidari et al., 2020; Simsek et al., 2020) also are very useful to use
for the identification of edible fig germplasm.
There are significant differences among the plant characteristics of caprifigs,
similar to edible figs (Table 7.2). Khadivi-Khub and Anjam (2014) reported that
caprifigs from Iran have numerous variables for tree shape and vigor, tree habit, the
number of lobes per leaf, leaf shape (Fig. 7.2), leaf length, leaf width, length of the
central lobe, little lateral lobe number, petiole length, and petiole thickness. Caliskan
et al. (2017b) showed that high diversity is determined in mamme and mammoni
number per shoot, ostiole width, terminal bud color, pollen number per fruit, fruit
7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 179

Table 7.1 The most diverse plant and leaf characteristics in the edible figs
Variable References
Beginning of fruit Mirheidari et al. (2020) and Simsek et al. (2020)
maturation
Cropping efficiency Simsek et al. (2020)
Tree growth habit Giraldo et al. (2010) and Pérez-Sánchez et al. (2016)
Size of the tree Giraldo et al. (2010) and Pérez-Sánchez et al. (2016)
Tree vigor Simsek et al. (2020)
Branching Mirheidari et al. (2020)
Apical dominancy Caliskan and Polat (2012a)
Degree of the Giraldo et al. (2010)
branching
Terminal bud length Gaaliche et al. (2012)
Shoot length Simsek et al. (2020)
Shoot color Mirheidari et al. (2020)
Leaf shape Saddoud et al. (2008), Caliskan and Polat (2012a), Baziar et al. (2018)
and Kokaj (2021)
Number of lobes Caliskan and Polat (2012a) and Rodrigues et al. (2019)
Shape of lobes Saddoud et al. (2008)
Upper lateral lobe Mirheidari et al. (2020)
depth
Upper lateral lobe Mirheidari et al. (2020)
base width
Leaf length Podgornik et al. (2010), Caliskan and Polat (2012a), Gaaliche et al.
(2012), Ciarmiello et al. (2015) and Kokaj (2021)
Leaf width Podgornik et al. (2010), Caliskan and Polat (2012a), Gaaliche et al.
(2012), Ciarmiello et al. (2015) and Kokaj (2021)
Leaf area Podgornik et al. (2010), Caliskan and Polat (2012a), Ciarmiello et al.
(2015), Kokaj (2021) and Khan et al. (2022)
Length of the central Caliskan and Polat (2012a) and Pérez-Sánchez et al. (2016)
lobe
Density of hairs on the Podgornik et al. (2010) and Mirheidari et al. (2020)
leaf
Petiole length Khan et al. (2022)
Petiole thickness Podgornik et al. (2010), Mirheidari et al. (2020)
The number of leaves Caliskan and Polat (2012a) and Simsek et al. (2020)
per shoot
The number of fruits Caliskan and Polat (2012a) and Simsek et al. (2020)
per shoot

skin color, internal pulp color, leaf shape, shoot length, and tree growth habit among
the caprifigs from Turkey. In addition, Mirheidari et al. (2020) indicated that ripen-
ing time, upper lateral lobe depth, and upper lateral lobe base width are important
variables for identifying caprifigs.
Generally, plant growth habit mainly has open and spreading habits; tree vigor is
most frequently classified as intermediate; shoot color is most often grey or brown;
180 O. Caliskan et al.

Table 7.2 The most diverse plant and leaf characteristics in the caprifigs
Variable References
Ripening time Mirheidari et al. (2020) and Khan et al. (2022)
Tree shape Khadivi-Khub and Anjam (2014) and Caliskan et al. (2017a)
Tree vigor Khadivi-Khub and Anjam (2014) and Caliskan et al. (2017a)
Tree habit Essid et al. (2017) and Mirheidari et al. (2020)
Shoot length Caliskan et al. (2017a)
Shoot color Caliskan et al. (2017b)
Terminal bud length Essid et al. (2017)
Terminal bud diameter Essid et al. (2017)
The number of lobes per Khadivi-Khub and Anjam (2014) and Caliskan et al. (2017b)
leaf
Leaf shape Khadivi-Khub and Anjam (2014), Caliskan et al. (2017b), Essid et al.
(2017) and Fatahi et al. (2017)
Leaf length Khadivi-Khub and Anjam (2014), Caliskan et al. (2017b), Essid et al.
(2017) and Mirheidari et al. (2020)
Leaf width Khadivi-Khub and Anjam (2014), Caliskan et al. (2017b), Essid et al.
(2017) and Mirheidari et al. (2020)
Leaf area Caliskan et al. (2017b)
Length of the central Khadivi-Khub and Anjam (2014), Essid et al. (2017) and Mirheidari
lobe et al. (2020)
Little lateral lobe Khadivi-Khub and Anjam (2014) and Mirheidari et al. (2020)
number
Upper lateral lobe depth Mirheidari et al. (2020)
Upper lateral lobe base Mirheidari et al. (2020)
width
Petiole length Khadivi-Khub and Anjam (2014)
Petiolethicknesss Khadivi-Khub and Anjam (2014)
Leaf number per shoot Caliskan et al. (2017a)

and the leaf area values of caprifigs are generally smaller than those of edible figs
(Caliskan et al., 2017b; Mirheidari et al., 2020). In addition, most caprifigs have five
or three lobes per leaf in Turkey (Caliskan et al., 2018), whereas it has mostly three
lobes per leaf in Iran (Khadivi-Khub & Anjam, 2014). This situation clearly shows
that the origin areas affect the difference among caprifig genetic resources. Besides,
Özen et al. (2017) indicated that the leaf length, width, area, petiole length, and thick-
ness of hybrid figs are lower than parents due to negative transgressive segregation.

3 Fruit Characteristics

Fig occurs in many genotypes, which have evolved mainly as natural seedlings dur-
ing the many centuries in fig-growing origin areas and in other continental countries
where it spread. Several cultivated and wild forms of fig are found in the origin areas
with a great diversity of color, shape, and flavor primarily for fresh consumption
7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 181

Fig. 7.2 Leaf shapes of figs: (a) Entire, (b) Base decurrent, (c) Base truncate, (d) base cordate, (e)
Base calcarate, lobes latete, (f) Base calcarate, lobes lyrate, (h) Base calcarate, lobes lineate

(Baziar et al., 2018; Caliskan & Polat, 2012a; Fatahi et al., 2017; Gozlekci, 2011;
Khadivi-Khub & Anjam, 2014; Sezen et al., 2014; Simsek et al., 2017). Fruit shape,
fruit size, small ostiole opening, ease of peeling (Aksoy et al., 2003; Condit, 1941),
total soluble solids content (TSS), TSS/acidity (Caliskan & Polat, 2012b; Crisosto
et al., 2010), flavor and fruit skin and flesh color (Ercisli et al., 2012; Flaishman
et al., 2008; Gozlekci et al., 2011) are evaluated as the main quality criteria of fresh
figs. Although these attributes are affected by ecology, they are mainly controlled by
the genetic capacity of the genotype (Caliskan & Polat, 2012b). Therefore, these
fruit quality characteristics are important in identifying fig genotypes. Distinct fruit
characteristics from these studies to date are summarized in Table 7.3.
The most important discriminators of edible figs are the fruit weight, fruit length,
fruit diameter, fruit shape (Giraldo et al., 2010; Podgornik et al., 2010), firmness of
the fruit skin (Podgornik et al., 2010; Saddoud et al., 2008), production type (Aljane
& Ferchichi, 2009; Podgornik et al., 2010), abscission of the stalk from the twig
(Podgornik et al., 2010), stalk diameter, neck diameter, ostiole diameter, ostiole
opening, and flesh thickness (Aljane & Ferchichi, 2009). In addition, crossing and
soma-clonal differences contribute to such diversity among fig genotypes (Ergül
et al., 2021).
Longitudinal cracks in the fruit peel and the ostiole opening (Fig. 7.3) are valu-
able features in identifying genotypes (Caliskan & Polat, 2012a; Saddoud et al.,
2008). Although accepted as a maturity criterion, the first is an undesirable feature
in the fresh fig trade. The second one, besides the genotype, may occur due to water
stress and air humidity, especially close to harvest.
182 O. Caliskan et al.

Table 7.3 The most diverse physical and chemical characteristics in the edible figs
Variable References
Production type Aljane and Ferchichi (2009) and Podgornik et al. (2010)
Abscission of the stalk Podgornik et al. (2010)
from the twig
Fruit shape Giraldo et al. (2010), Ciarmiello et al. (2015) and Baziar et al. (2018)
Fruit weight Podgornik et al. (2010), Caliskan and Polat (2012a), Fatahi et al.,
2017; Abdelsalam et al. (2019), Hssaini et al. (2020a), Simsek et al.
(2020)
Fruit diameter Saddoud et al. (2008), Pérez-Sánchez et al. (2016), Fatahi et al., 2017;
Hssaini et al. (2020a) and Simsek et al. (2020)
Fruit length Podgornik et al. (2010), Pérez-Sánchez et al. (2016), Fatahi et al.,
2017; Hssaini et al. (2020a), Simsek et al. (2020) and Khan et al.
(2022)
Fruit flesh thickness Mirheidari et al. (2020)
Fruit neck length Podgornik et al. (2010), Ciarmiello et al. (2015), Pérez-Sánchez et al.
(2016), Hssaini et al. (2020a) and Simsek et al. (2020)
Ostiole opening Aljane and Ferchichi (2009)
Ostiole width Aljane and Ferchichi (2009) and Gaaliche et al. (2012)
Fruit skin color Saddoud et al. (2008), Caliskan and Polat (2011), Gozlekci (2011) and
Hssaini et al. (2020a)
Fruit flesh color Saddoud et al. (2008)
Firmness of the fruit Saddoud et al. (2008) and Podgornik et al. (2010)
skin
Fruit skin thickness Saddoud et al. (2008)
Fruit skin cracks Saddoud et al. (2008) and Caliskan and Polat (2012a)
Easy of peeling Ciarmiello et al. (2015) and Pérez-Sánchez et al. (2016)
Drop at the eye Simsek et al. (2020)
Color of liquid drop at Simsek et al. (2020)
the ostiole
Total soluble solids Giraldo et al. (2010), Caliskan and Polat (2012a), Ciarmiello et al.
(TSS) (2015), Pérez-Sánchez et al. (2016) and Hssaini et al. (2020a)
pH Simsek et al. (2020)
Acidity Messaouidi and Haddadi (2008), Gaaliche et al. (2012) and Simsek
et al. (2020)
TSS/acidity Caliskan and Polat (2012a) and Simsek et al. (2020)
Total antioxidant Caliskan and Polat (2012a)
capacity
Total phenolic Caliskan and Polat (2012a) and Hssaini et al. (2020b)
Total anthocyanin Caliskan and Polat (2012a) and Hssaini et al. (2020a)
Glucose Gozlekci (2011), Caliskan and Polat (2012a) and Hssaini et al. (2020b)
Fructose Gozlekci (2011), Caliskan and Polat (2012a) and Hssaini et al. (2020b)

Fruit skin and flesh color of fresh figs are very important for consumer prefer-
ences. Consumers prefer fresh figs with dark skin color and red or pink flesh color.
However, the main crops’ most frequent fruit skin color is green, and the pulp is red
and pink (Caliskan & Polat, 2012a; Condit, 1941; Giraldo et al., 2010). In addition,
7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 183

Fig. 7.3 Classification of cracking in fig fruit peel (upper) and ostiole width (lower)

Flaishman et al. (2017) reported that the black color of the skin is dominant over the
green and red flesh color is dominant over pale color. Moreover, figs are one of the
richest fruits in terms of skin color and flesh color diversity. According to the fig
descriptor, there are seven different fig skin colors and five fruit flesh color (Fig. 7.4)
groupings. This rich color difference in figs constitutes one of the essential criteria
for identifying fig genotypes (Caliskan & Polat, 2011; Gozlekci, 2011; Hssaini
et al., 2020a; Saddoud et al., 2008). Also, the skin coloration of figs is not homoge-
neous, especially in colored genotypes, which can cause differences in color evalu-
ation such as green-brown and purple-black. However, the more objective conversion
of these color observations into unbiased values results in more successful discrimi-
nation of various genotypes. For this purpose, using a colorimeter instead of a color
chart can be preferred in descriptive and selective research to obtain more reproduc-
ible results. Indeed, Gozlekci (2011) indicated that fig genotypes’ lightness (L),
chroma, and hue° values of skin color vary greatly. Also, Caliskan and Polat (2011)
reported that for figs with a vast diversity of fruit color among genotypes, objective
investigations were very appropriate due to the researcher’s subjective observation
changes.
In describing the genetic sources of figs, the physical measurements and phyto-
chemical properties can be used as suitable criteria. Fig cultivars with dark skin
contain higher levels of polyphenols, anthocyanins, and flavonoids accompanied by
higher antioxidant activity than fig genotypes with lighter skin (Aljane et al., 2020;
Caliskan & Polat, 2011; Solomon et al., 2006). Caliskan and Polat (2011) reported
that the dark genotypes contained the highest amounts of total antioxidant capacity,
total phenolics, and total anthocyanins, whereas the yellow and green genotypes
contained the lowest amounts. Previous studies on figs have focused on selecting
and describing plant characteristics, fruit quality characteristics, and genetic mark-
ers. In recent years; however, because of the increasingly apparent importance of a
healthy diet, researchers may also need to include phytochemical analysis in germ-
plasm evaluation. The total antioxidant capacity, total phenolics, anthocyanins,
fructose, and glucose contents of the fig fruits are; besides, Caliskan and Polat
(2012b) showed that the sugar content of figs negatively correlated with total
184 O. Caliskan et al.

Fig. 7.4 Fruit skin and flesh color grouping in edible figs. Fruit skin color groups: 1 Black, 2
Purple, 3 Brown, 4 Green, 5 Light green, 6 Yellow-green, 7 yellow. Fruit flesh groups: 1 White, 2
Amber, 3 Pink, 4 Red, 5 Dark red

anthocyanins due to the sugar content of the cultivars with yellow and green fruit
skin color being higher than those with dark colors.
Caprification is a common application in fig-growing areas and significantly
affects the fruit set in edible figs (Condit, 1947). Caprifigs have a positive effect not
only on fruit set and yield but also on quality characteristics such as fruit size, fruit
flesh color, TSS content (Gaaliche et al., 2011; Ferrara et al., 2016), aroma and
phytochemical content (Pourghayoumi et al., 2012; Trad et al., 2012) of edible figs.
Important variables of fruit characteristics in the caprifigs are shown in Table 7.4.
Acarsoy Bilgin et al. (2020) indicated that identifying caprifig genetic material is
important for caprification and breeding studies. Unfortunately, there have been a
few studies into the genetic resources possible for caprifig germplasm. Further, cap-
rifig accessions’ morphological and pollinizer parameters have not been detailed.
On the other hand, all caprifigs are not helpful in caprification because of important
parameters in their unique fruit characteristics.
The critical characteristics of the caprifig for profichi crops (Table 7.4) include
fruit size, fruit number per shoot, the time of Blastophaga wasps exit from caprifig
fruits, ripening period, coinciding with the female flowers in edible figs (Caliskan &
Bayazit, 2012), amount of gall, and male flowers, amount of pollen production (Ahi
Koşar et al., 2022; Yaman & Caliskan, 2016a), pollen viability and germination
ratios (Ilgin et al., 2007; Yaman & Caliskan, 2016b), the inclusion of the mammoni
7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 185

Table 7.4 Important variables of fruit characteristics in the caprifigs

Variable References
Fruit shape Caliskan et al. (2017b)
Fruit weight Khadivi-Khub and Anjam (2014), Caliskan et al. (2017b), Essid
et al. (2017) and Acarsoy Bilgin et al. (2020)
Fruit width Khadivi-Khub and Anjam (2014), Essid et al. (2017) and
Acarsoy Bilgin et al. (2020)
Fruit length Caliskan et al. (2017b) and Acarsoy Bilgin et al. (2020)
Fruit neck length Caliskan et al. (2017b)
Fruit stalk length Khadivi-Khub and Anjam (2014) and Essid et al. (2017)
Ostiole width Caliskan et al. (2017a) and Essid et al. (2017)
Fruit internal pulp width Khadivi-Khub and Anjam (2014)
Fruit skin color Khadivi-Khub and Anjam (2014) and Caliskan et al. (2017b)
Fruit internal color Caliskan et al. (2017b)
Profichi: Date of Blastophaga Caliskan et al. (2017b)
emergence
Profichi: Duration of Caliskan et al. (2017b)
Blastophaga emergence
Profichi: The number of gall Caliskan et al. (2017b)
flowers
Mammoni number per shoot Caliskan et al. (2017a)
Mamme number per shoot Caliskan et al. (2017a)
The numbers of pollen grains Caliskan et al. (2017b)
per fruit
The numbers of pollen grains Caliskan et al. (2017b)
per flower
The numbers of pollen grains Caliskan et al. (2017b)
per anther
Pollen viability Caliskan et al. (2017b)
Pollen germination Caliskan et al. (2017b)
Pollen shape Caliskan et al. (2021)
Pollen diameter Acarsoy Bilgin et al. (2020) and Caliskan et al. (2021)
Pollen length Acarsoy Bilgin et al. (2020) and Caliskan et al. (2021)
Colpus width Caliskan et al. (2021)
Exine thickness Caliskan et al. (2021)
Number of single porates Caliskan et al. (2021)
Number of double porates Caliskan et al. (2021)
Number of triple porates Caliskan et al. (2021)
Porate width Acarsoy Bilgin et al. (2020) and Caliskan et al. (2021)
Abnormal pollen ratio Caliskan et al. (2021)

and mamme crops, and free from disease and pests (Caliskan et al., 2016; Yaman &
Caliskan, 2016a).
The fruit shapes of the caprifigs are mostly oblong, globose, or oblate. After pol-
lination, the flowers within fruits of edible figs may fill the syconium cavity as they
mature and form a solid pulp that constitutes about 83% of the weight of a mature
186 O. Caliskan et al.

fig. However, this does not occur in caprifigs (Condit, 1947), which have low spe-
cific gravity due to the large intercellular air spaces in the fruit’s peel compared to
edible figs (Neeman & Galil, 1978). Thus, the fruit weights, particularly caprifigs,
can be lower than edible figs. The caprifigs have green skin color and white internal
color. In addition, Caliskan et al. (2017b) reported that mamme and mammoni crops
could be varied by genotypes. All caprifigs have a profichi crop set; however, the
numbers of fruit differed depending on genetic and ecological conditions. Also, the
profichi fruit yield is higher than that of edible figs.
Khadivi-Khub and Anjam (2014) showed that fruit and leaf size, fruit shape, leaf
shape, and fruit skin color are important parameters for the morphological charac-
terization of caprifigs. Caliskan et al. (2017b) stated that the fruit skin color of male
figs is not as rich as that of female figs and that 90% of the genotypes are green in
color, and there are a small number of black, purple, and brown skin colored geno-
types (Fig. 7.5). Besides, the internal pulp color of caprifigs is most often white,
whereas a color distribution ranging from dark purple to light purple can be seen in
the fruit pulp color (Fig. 7.6).
Caliskan et al. (2017b) reported that the number of gall flowers per fruit, date of
Blastophaga emergence, duration of Blastophaga emergence, percentages of pollen
viability and germination, the number of pollen grains per anther, the number of
pollen grains per flower, and the number of pollen grains per fruit that could be suc-
cessfully used to characterize caprifig accessions. Besides, caprifig classes accord-
ing to the number of gall flowers per fruit are mostly medium (250–499) and high
(500–749), and the number of male flowers per profichi fruit ranges between 50 and
150 (Caliskan et al., 2018). They revealed great genetic diversity and intensive

Fig. 7.5 Fruit skin color differences in caprifigs

7 Morpho-Chemical Characteristics Useful in the Identification of Fig (Ficus… 187

Fig. 7.6 Fruit internal

color differences in
caprifigs

differentiation of caprifigs. This rich genetic variation could have been due to the
establishment of caprifig populations propagated by seed.
Caliskan et al. (2017b) reported a few morphological characteristics such as the
fruit size, the number of gall flowers per fruit, pollen viability and germination, and
pollen number per anther that could be used for the differentiation of caprifigs.
Thus, more distinctive variables are essential to define germplasm and establish a
breeding program for caprifigs.
Pollen grains have specific morphological characteristics and demonstrate pow-
erful genetic preservation; therefore, they can be used in taxonomy, phylogeny,
morphological characterization, and breeding studies of fruit species (Shivanna,
2003; Arzani et al., 2005; Evrenosoglu & Misirli, 2009). Caliskan et al. (2021)
showed that polar length, equatorial diameter, colpus width, pollen shape, number
of porates (Fig. 7.7), porate width, exine thickness, and abnormal pollen ratio are
the most important properties in distinguishing caprifigs from each other. Therefore,
these pollen characteristics can also be used to identify caprifigs.

4 Conclusions

Selecting highly distinctive variables for appropriate morphological characteriza-

tion of fruit genetic resources is invaluable for sustainably optimizing resources.
This should be necessary for fig germplasm studies where many similar,
188 O. Caliskan et al.

Fig. 7.7 Porate number of caprifig pollens: Single-porate (left), double-porate (middle), and
triple-porate (right)

homonymous, and synonymous genotypes can be observed. Some morpho-chemi-

cal characters show significant variability in edible figs and caprifigs. Plant charac-
teristics such as ripening time and leaf characteristics, fruit characteristics such as
size, shape, skin and ostiole cracking, skin and flesh color, and chemical properties
such as TSS, acidity, TSS/acidity, sugar profile, and phytochemical contents were
found to be very successful in distinguishing genotypes from each other in edible
figs. These characteristics can be used together to identify genotypes and the selec-
tion of individuals before breeding. In addition, fruit characteristics such as fruit
size, skin and pulp color, the number of gall flowers, pollen number per fruit, and
pollen characteristics such as pollen size and shape and pore number are successful
in distinguishing individuals in a profichi crop of caprifigs. This review will be an
important contribution to selecting highly discriminative variables to optimize
resources to evaluate fig germplasm.

References

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Chapter 8
Agronomic Strategies for Fig Cultivation
in a Temperate-Humid Climate Zone

Norma Micheloud, Paola Gabriel, Juan Carlos Favaro, and Norberto Gariglio

1 Introduction

The fig (Ficus carica L.), apparently the earliest known cultivated fruit crop (Kislev
et al., 2006), is a small deciduous tree native to Western Asia, which has been dis-
tributed and cultivated throughout the Mediterranean region, being Turkey, Egypt,
Iran, Greece, Algeria, Morocco, Syria, Italy, and Spain the main producing coun-
tries (Flaishman et al., 2008). In the northern hemisphere, it is cultivated between 35
and 40° S. In the American continent, its cultivation is widespread mainly in the
United States and Brazil (Nieto et al., 2007), Peru, Bolivia, and Argentina
(Morton, 2013).
The species of the genus Ficus have fruits of little commercial interest, except for
the fig (Sánchez de Lorenzo, 2009), which is considered one of the healthiest fruits
associated with longevity (Trichopoulou et al., 2007) and food security (Barolo
et al., 2014). In addition, this fruit plays an essential role in human nutrition due to
its high vitamin, mineral, and fiber content (Venu et al., 2005). Although figs can be
used for direct consumption as fresh fruits, since they are highly perishable even in
refrigerated conditions (Piga et al., 1995), they are preferentially eaten dried as the
main or auxiliary ingredient in many desserts (Uzundumlu et al., 2018) or as fruit
powder (Khapre et al., 2015). Figs are also considered excellent for processing fruit
juice and fruit tea (Lim, 2016). Dried figs can also be mixed with nuts for snacks or

N. Micheloud · N. Gariglio (*)

ICiAgro Litoral, UNL, CONICET, FCA, Esperanza, Santa Fe, Argentina
e-mail: [email protected]; [email protected]
P. Gabriel · J. C. Favaro
Facultad de Ciencias Agrarias, Universidad Nacional del Litoral,
Esperanza, Santa Fe, Argentina
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 193
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_8
194 N. Micheloud et al.

cereals for muesli, while dried fig puree can be used as a raw material for ice cream
and marmalades (Aksoy, 2017).
Although the fig may be the oldest cultivated fruit species, little is known about
its production practices, and the low number of available cultivars limits its global
expansion (Botti et al., 2003). However, it is worth noting that figs are relatively
easy to produce in an agro-ecological way (López-Corrales & Balas, 2014) and that
the fig has excellent commercial potential due to its high market prices, the increases
in demand, and the health benefits of its fresh and dried fruits (Abbas et al., 2019;
López-Corrales & Balas, 2014).

2 Eco-physiological Requirements of Fig Tree

The fig is a functionally dioecious species with numerous small flowers and fruits
located inside a fleshy receptacle called syconium (López-Corrales & Balas, 2014).
Fig growth and production are strongly dependent on climatic conditions. Generally,
the fig grows best and produces high-quality fruit in the Mediterranean and dry
warm-temperate climates, with hot summers and mild winters, low rainfall, and
relative humidity (Gaaliche et al., 2011). To express maximum fruit yield, the fig
tree also requires high radiation. However, fig trees can also grow adequately under
less favorable conditions (Leonel & Tecchio, 2010), although crop production is
affected by the decrease in temperature during autumn, the cold winter conditions,
and rainfall mainly during fruit maturation (Flaishman et al., 2008). Under colder
weather, the tree stops growth, becomes defoliated, develops a typical terminal bud,
and enters a dormancy period (Kawamata et al., 2002).
The fig tree has low chilling requirements to break dormancy and tolerates light
frost (Limeira Da Silva et al., 2016). However, low winter temperatures constitute a
limiting factor for its commercial cultivation, being −12.2 °C is the threshold of tree
death by frost, although young trees may be damaged by higher frost temperatures
between −5 and −10 °C (Ferguson et al., 1990). Tolerance to cold dramatically
depends on the cultivar‘s origin (López & Salazar, 2010).
In contrast, despite its deciduous characteristics, under hot climatic conditions
such as those in Brazil, the tree can grow continuously and behave as evergreen
because with night temperatures above 12 °C, the plant continues to grow, and
fruits ripen.
The climatic conditions also affect the crop type (Breba or Main), the harvesting
time, and the fruit quality. For example, in cultivars sensitive to frost and/or low
winter temperature conditions, early or late frosts kill the terminal part of the cur-
rent year shoots, affecting the Breba crop (Ferguson et al., 1990); consequently,
Breba production is successful only under relatively moderate winters.
The annual fig growth and phenology also respond to thermal accumulation,
calculated by considering a minimum basal temperature of 8 °C and a maximum
temperature of 36 °C (Souza et al., 2009). For example, in the municipality of
Botucatu, in the state of São Paulo, Brazil, Ferraz et al. (2020) found that the cv.
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 195

‘Roxo de Valinhos’ requires 167 days or 2458 degree-days from pruning to the
beginning of the harvest period. Temperature conditions also affect the color and
shape of the fruits. In colder climates, the pulp takes on more intense color, and the
fruits are more elongated; and since the fruits do not suffer damage by cooling, they
can be stored at up to −1 °C and 90–95% of relative humidity (Caliskan & Polat,
2011). On the other hand, a comparison between dried and fresh figs has shown that
dried fig production is highly dependent on climatic conditions, successful in dry
and warm temperate climates. In contrast, fresh figs can grow in broader ecological
conditions (López-Corrales et al., 2011; Sahin, 1998).
Fig trees adapt well to various soil types, except those with poor drainage
(Flaishman et al., 2008). They can be grown in heavy clays, barns, and light sands,
but ideally, the soil should be well-drained and with good fertility. Fig trees are
tolerant to high soil calcium content (Aksoy, 1998) and are among the few fruit trees
of temperate zones with moderate resistance to salinity (Ayers & Westcost, 1985).
Upon root exposure to 100 mM sodium chloride (NaCl), fig leaves remain healthy
and green, and gas exchange parameters also remain relatively high (Caruso et al.,
2017). Their resistance to salinity is comparable to that exhibited by olive plants
(Vangelisti et al., 2019). Fig genes may thus be useful as targets for gene editing to
increase salt tolerance in figs or other tree species (Vangelisti et al., 2019).
Fig trees are also resistant to drought and perform well in semi-arid climate
regions. Although rainfed cultivation is widely spread, the growth and productivity
of fig trees respond positively to irrigation (Tapia et al., 2003). Furthermore, the
extensive root system of the fig tree allows it to be used for both soil restoration and
erosion control, as well as for rainforest regeneration because of the seed rain, seed
bank, and vegetation established beneath the canopies of fig trees (Guevara
et al., 2004).
In fig plantations, rootstocks are not usually used because the commercial variety
is usually propagated by cuttings (López-Corrales & Balas, 2014). Wood cuttings
are taken from one-year-old shoots, and rooting is not a problem for growers; the
main adversity is the loss of plants by soil pathogens and nematodes (Kilinç et al.,
2007). Therefore, soilless cultivation in nursery production as one of the safest alter-
natives to reduce plant loss has expanded enormously since the 1980s (Chong &
Lumis, 2000; Evans et al., 1996; Şirin et al., 2010), and studies have been conducted
to adapt the use of widely available regional substrates for fig cultivation (Wright
et al., 2006).

3 Climatic Conditions of the Central Area of Santa Fe

and Their Suitability for Fig Cultivation

The fig is mainly cultivated in semiarid regions (Gaaliche et al., 2011; Limeira Da
Silva et al., 2016). Thus, in Argentina, the main fig cultivation areas are located in
the country’s northwest, in the Cuyo region, and in Buenos Aires province
196 N. Micheloud et al.

(Prataviera, 2003), with the predominance of three varieties: Brown Turkey,

Guarinta, and Kadota. Since the production is not enough to supply the domestic
market, dried figs are imported from Turkey and Chile annually. In addition, new
plantations in Argentina have been adapted to high tree densities (>1000 plants per
hectare), and the canopy is annually renewed by pruning (Prataviera, 2003), which
allows trees to remain small, facilitating the horticultural practices.
Fig crops have also been spread as a complementary commercial activity of tra-
ditional horticultural crops in the horticultural area around Santa Fe city (31°26′ S;
60°56′ W), in the central-east region of Argentina (Travadelo et al., 2017), although
the climate of this area is temperate humid mesothermal (Fig. 8.1) classified as Cfa
(Köppen, 1948).
In this region, the mean annual global radiation is 16 MJ m−2 day−1, with maxi-
mum and minimum values of 23 and 9 MJ m−2 day−1 for January and July, respec-
tively. In the summer and winter solstice, the astronomical heliophany ranges
between 14 and 10 h, respectively (García et al., 2014). The average temperatures of
the warmest (January) and coolest months (July) are 26.7 °C and 11.7 °C, respec-
tively (García et al., 2014). The absolute minimum temperature reaches an average
value of −3.9 °C, and frosts occur between May and September (García et al.,
2013). The availability of cold has an average value of 301 chilling hours, with a
high coefficient of variation (31%) (Gariglio et al., 2012).
Precipitations have an irregular distribution, with a tendency to the Monsoon
pattern (Papadakis, 1952). The annual precipitation is near 1200 mm on average.
The minimum rainfall values occur during the winter (June, July, and August) with
a monthly average value of 23 mm. Precipitations increase in spring and reach their

900 35

800
30
700
25
Temperature (°C)
Rad (Mj m-2)

600
Pp (mm)

500 20

400 15
300
10
200
5
100

0 0
J F M A M J J A S O N D

Precipitation (mm) Radiation (Mj m-2) T MAX (°C) T MIN (°C)

Fig. 8.1 Monthly mean values of accumulated solar radiation (Rad) and precipitation (Pp), maxi-
mum temperature (TMAX), and minimum temperature (TMIN) recorded in the central area of
Santa Fe province (Argentina) during 2007–2016. (Adapted from Micheloud et al., 2018)
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 197

maximum during the summer, with a monthly average of 164 mm. The annual num-
ber of rainy days is 92 ± 10 days, being March and November the months with the
highest value, and floods usually occur during February and March. The relative
humidity is higher during summer, ranging between 72 and 80%, with an annual
value of 76% (García et al., 2014).
The soils have subnormal to normal relief, and it is common to find vast plains
with slopes less than 1%. The predominant soils are typical Argiudolls, aquic
Argiudolls, and Argialbolls; the best soils in this area show a slight limitation due to
the clayey texture of the subsoil (B horizon). On the other hand, the soils near the
Paraná River are sandy, with slight to moderate limitations due to their low water
retention capacity; however, they are very suitable for horticultural crops with drip
irrigation (García et al., 2014).

4 Climatic Conditions with Adverse Effects of Humidity

on the Fig Tree and Its Fruits

Figs are climacteric fruits slightly sensitive to the action of ethylene on promoting
softening and decay, especially when fruits are kept over 5 °C (Gözlekçi et al.,
2008). Since fruit maturation induces important structural and chemical changes the
harvesting time is a determining factor of fruit quality attributes, mainly flavor, with
the particularity that overripe fruits can become undesirable due to fermentative
products (Martínez-González et al., 2017; Owino et al., 2006). The manifestation of
these changes depends on the cultivar and the agro-ecological conditions (López-
Corrales et al., 2011; Morton, 2013).
Fruit quality is also affected by high humidity and frequent rains; mainly if ripen-
ing occurs under these unfavorable conditions, in which case, fruits develop vine-
gary odors that affect their commercialization (Nienow et al., 2006; Sala, 2000).
Furthermore, fig fruits are prone to cracking or splitting at maturity due to rains or
abundant irrigation (Ferguson et al., 1990; Nienow et al., 2006) (Fig. 8.2). Fruit
quality and production can also be reduced by other environmental conditions such
as hail and wind (Martínez-González et al., 2017).
Very little research has been done to extend the postharvest life of fresh figs;
however, low temperatures (0–2 °C) and high relative humidity (90–95%) are rec-
ommended. Fruit deterioration occurs mainly by the incidence of microbial molds
and rots (Cantín et al., 2011). It has been found that figs kept at 2 °C with modified
atmosphere packaging can be stored for 21 days and have better internal and exter-
nal appearance than control fruits, which show significant loss of quality by dehy-
dration (18%) (Bouzo et al., 2012).
To avoid water logging in the base of the trunk and principal roots during abun-
dant rainfall periods and possible plant flooding damage, fig trees are cultivated on
20-cm-high ridges (Fig. 8.3). Low oxygen supply to the roots caused by soil
198 N. Micheloud et al.

Fig. 8.2 Fig fruits affected by ‘splitting’ caused by excess water during maturation in Santa Fe
(Argentina)

flooding greatly reduces the energy status of root cells, limiting the growth, devel-
opment, and survival of fig trees (An et al., 2016).

5 Alternatives to Reduce the Adverse Effects of Rains

During the Fruit Ripening Period

Green figs are hard, have a rubbery feel, and cannot ripen off the tree; however, fruit
picked just before full ripeness will continue to soften and become soft to the touch
and sweet at room temperature, and the riper the fig, the sweeter it is. Consequently,
the adverse effects of rains can be diminished by harvesting the fruits for the fresh
market just before full ripeness. Accordingly, at the time of full harvest and high-
temperature conditions, as it occurs during summer in the central area of Santa Fe,
harvesting must be carried out daily (January and February in the southern hemi-
sphere), while when the temperature decreases toward the end of summer and
autumn (March to May), the frequency between harvests may decrease up to a once
a week (Gariglio et al., 2014). Therefore, the organization of this cultural practice
and the estimation of its labor demand are crucial aspects of the fig crop because of
its perishable fruit and long harvesting period.
The application of ethylene to advance fig fruit ripening proves suitable (Owino
et al., 2006). Ethylene is considered the plant hormone responsible for fruit
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 199

Fig. 8.3 Fig trees, cv.

‘Brown Turkey’ cultivated
on raised ridges to avoid
excess water accumulation
in the base of the trunk due
to the abundant rainfall
that commonly occurs
during autumn in Santa Fe
(Argentina)

ripening, and some of its physiological effects include the modification of fruit
color due to the degradation of chlorophylls and the synthesis of carotenoids and
flavonoids; the modification of the fruit texture by changes in the cell turgor and the
cell wall structure; the modification of the concentration of sugars, organic acids
and volatile compounds that affect the fruit nutritional quality, flavor and aroma;
and the increase in the susceptibility to be attacked by opportunistic pathogens that
are associated with the loss of integrity of the cell wall (Martínez-González
et al., 2017).
Applying ethylene to fig fruits during the late stage of their development acceler-
ates the ripening process by stimulating endogenous ethylene synthesis. As a result,
treated fruits reach a greater size, have higher soluble solid content, and have better
coloration, texture, and flavor, being able to be harvested 7–14 days after the appli-
cation according to the temperature (Melgarejo-Moreno, 1999). Because of the fast
effect of ethylene on fig fruit ripening, it is possible to advance fruit harvesting
when a prolonged rainy period is predicted and consequently reduce the adverse
effect on fruit quality. The treatment must be directed only to the fruits that are
going to be harvested in that period (unpublished data).
200 N. Micheloud et al.

6 The Extension of the Harvest Period as an Alternative

to Reduce the Risk of Fruit Damage by Rains During
fruit Maturation

In areas where rainfall does not have a seasonal distribution or shows high variabil-
ity between years, as in the central area of Santa Fe (Argentina), an alternative to
reduce the risk of fruit damage by rainfall during ripening is the extension of the
harvesting period. A period of excessive rains in coincidence with a short harvest
period can damage a greater proportion of the annual fruit production. In contrast,
only the proportion of fruits that ripen during the rainy period will be damaged in a
long harvesting strategy. Furthermore, a long harvesting period is favorable for
fresh market supply. Since figs ripen sequentially along with the shoot, picking
should be done repeatedly during harvest (Flaishman et al., 2008).

6.1 Promotion of Stem Growth as a Strategy to Extend

the Harvest Period

Fig cultivars may produce one or two crops a year: the Breba crop and the Main
crop. The fruit of the Breba crop is born laterally along with the shoot of the previ-
ous season, whereas the fruit of the Main crop is produced laterally along with the
shoot of the current season (Flaishman et al., 2008). As a consequence of the axil-
lary fruit location, fig yield strongly depends on the vegetative growth, so the num-
ber of fruits of each crop is closely related to shoot length (Gaaliche et al., 2011).
The Breba crop depends on the length of one-year-old shoots, whereas the Main
crop depends on the length of the shoot of the current season; furthermore, the Main
crop indirectly affects the Breba crop for the next year (Gaaliche et al., 2011). In the
warm climate region of São Paulo, Brazil, for example, since figs of the Main crop
ripen sequentially along with the current shoot, longer shoots cause the extension of
the harvesting period (Leonel & Tecchio, 2010). Therefore, regular and high fig tree
productivity requires the promotion of shoot growth, which can be achieved by
applying effective cultural practices, such as irrigation, fertilization, and pruning.

6.1.1 Irrigation

As mentioned above, fig trees adapt to different soils, being the most appropriate
those of clayey-sandy texture and rich in organic matter (Pereira, 1981). Since the
fig is considered a rustic plant (Rodrigues et al., 2012), the largest traditional culti-
vation areas in the world are located in Mediterranean arid regions under rainfed
conditions (Çalişkan & Polat, 2011).
Despite its rusticity, excessively dry soil causes plants to remain resting, devel-
oping few leaves without producing fruits (Pereira, 1981). However, the fig is
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 201

currently one of the fruit species with great global expansion and a favorable
response to technology incorporation (Mendoza-Castillo et al., 2017). Under this
context, irrigation is an important technology for achieving high yield and fruit
quality (Melgarejo et al., 2007). For example, in the state of São Paulo, Brazil, irri-
gation of fig trees with a water supply of 75% of the Class A pan evaporation showed
positive effects on fruit yield, branch length, and fruit length and diameter
(Hernández et al., 1994). However, the irrigation frequency can also influence the
water requirement; for example, for 2- and 4-day irrigation intervals, the adequate
water supply can be reduced from a mean crop coefficient of 0.71 to 0.51, respec-
tively (Andrade et al., 2014).
In combination with severe pruning (see below), irrigation increases the diameter
of the current shoots, the number and length of internodes, and consequently, the
length of shoots (Norberto et al., 1998). A good positive correlation between the
irrigation rate and shoot length has also been observed in fig crops in the semi-arid
north-central zone of Chile (Tapia et al., 2003).

6.1.2 Nitrogen Fertilization

The literature concerning fig tree fertilization is scarce. The N:P:K ratio commonly
used in fig tree nutrition is approximately 20:5:20 (Flaishman et al., 2008). Under
the traditional cultivation system for dry figs in Greece and very calcareous, clayey,
and alkaline soils, Sotiropoulos et al. (2020) found that the treatment that resulted
in the highest yield was the single annual application of 1 kg of N, 1.2 kg of P and
0.6 kg of K per tree, during February. These authors found that, despite the adequate
fertilization with N, the trees showed a leaf N content lower than the critical value
of 1.7% d.w. In contrast, leaf K concentrations were found to be sufficient (K > 1%
d.w.), and leaf P content generally showed a marginal sufficiency (P > 0.1% d.w.)
(Sotiropoulos et al., 2020). In another experiment under hydroponics and green-
house conditions in Mexico, Mendoza-Castillo et al. (2019) found that the total dry
matter accumulation 135 days after pruning was 12.7 t ha−1, with a mineral extrac-
tion of 250 kg ha−1 of N, 80 kg ha−1 of P and 110 kg ha−1 of K. In this experiment,
the authors generated the curves of dry matter accumulation and mineral extraction,
which are essential to establish fertilization programs based on the extraction curves
and crop phenology.

6.1.3 Pruning Intensity

Traditionally, the fig tree was trained in an open-vase shape with three main branches
to obtain a tree less than 3 m in height (Melgarejo-Moreno, 1999). However, tree
height could be reduced, and plant density could be increased by minimal pruning,
which consists of the removal and/or head back of some branches (30%) to prevent
the basal and internal canopy senescence and allow tree rejuvenation and shoot
renovation (Lodolini et al., 2021).
202 N. Micheloud et al.

In different fruit tree crops, severe pruning increases the vigor of vegetative
shoots and reduces the number of flower buds (Greene, 1999). This effect is due to
the production of large amounts of gibberellins in young leaves and apical meri-
stems (Jones & Phillips, 1966), which inhibit flower induction (Cleland, 1969) and
reduce the flowering intensity the following year. In the fig winter pruning stimu-
lates the growth of the shoot, which is monopodial with the apical meristem devel-
oping leaves and inflorescences along the axis (Alvarenga-Gonçalves et al., 2006;
Gaaliche et al., 2011; Lodolini et al., 2021). However, shoot growth stimulated by
pruning does not affect fruit production of fig, as observed in other fruit trees; in the
fig, greater shoot growth should favor greater fruit production per shoot and a longer
harvesting period (Alvarenga-Gonçalves et al., 2006; Micheloud et al., 2018). This
behavior tends to spread the increases in pruning intensity of the fig tree, even with
an annual renewal of the canopy. In situations where this pruning and training sys-
tem is adopted, it has been associated with small plant size and high plant density.
Furthermore, this system has been adapted to greenhouse cultivation, allowing its
large expansion in different countries such as Mexico (Mendoza-Castillo et al.,
2017), Japan (Hosomi et al., 2015), and Malaysia (Shamin-Shazwan et al., 2019). In
fig orchards under rainfed conditions in Iran, severe pruning intensity has also been
long-term beneficial effects in reducing drought damages (Zare, 2021).

6.1.4 Other Strategies to Improve Stem Growth

Regarding the cold requirements of the fig tree, there are certain contradictions. Its
buds require little or no winter chilling to break endodormancy (Ferguson et al.,
1990). However, the evolution of dormancy in different cultivars under Moroccan
conditions studied using the biological test (single node cuttings) showed that the
removal of dormancy required the accumulation of 450 and 400 chill hours during
the first and second year of study, respectively (Oukabli & Mekaoui, 2012). This
cold requirement justifies the application of hydrogen cyanamide in warm areas to
increase fig bud break and stimulate earlier bud break and harvest of the Breba crop
(Gerber et al., 2010; Yablowitz et al., 1998). In Tunisia, the main fig cultivar of the
Smyrna type grown commercially is ‘Zidi’, a cultivar that produces only one Main
crop (Gaaliche et al., 2017). By studying this cultivar, Gaaliche et al. (2017) found
that hydrogen cyanamide (1.5%) advanced bud break by 10 days and increased
shoot length, fruit number per shoot, and yield. They also found that fruit ripening
occurred 7 days earlier than in controls. These authors thus concluded that, in this
warm area, hydrogen cyanamide could be used to improve vegetative growth and fig
productivity and promotes early fruit maturity, allowing the grower to take advan-
tage of the premium prices of the early season (Gaaliche et al., 2017).
According to the previous evidence, adopting the pruning system that causes the
annual canopy renewal of the fig tree requires ensuring adequate shoot growth since
the number of nodes and reproductive buds is related to shoot length (Alvarenga-
Gonçalves et al., 2006; Gaaliche et al., 2011). The longer the current shoots, the
greater the potential number of fruits per shoot and the longer the harvesting period.
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 203

Therefore, although the fig tree is a rustic plant that grows in arid areas, changing
the training system to annual canopy renewal requires good agronomic practices
and technological application.

7 Effects of Annual Renewal of the Canopy

As previously explained, the annual renewal of the canopy by pruning is a simple

strategy to promote shoot growth and extend the fig‘s harvesting period. Under the
traditional pruning and training system of the Mediterranean area and rainfed condi-
tions, the current shoots of the fig tree have an annual growth of about 10 cm
(Ateyyeh & Sadder, 2006), and trees can produce two crops a year (Breba and
Main) depending on the cultivar (Gaaliche et al., 2011). In the temperate climate
conditions of the central area of Argentina, in both the ‘Brown Turkey’ and
‘Guarinta’ cultivars, the annual renewal of the canopy causes the loss of the Breba
crop and dramatically increases the current shoot length over 139 cm, without
affecting the fruit set (Micheloud et al., 2018). Under the tropical conditions of
Minas Gerais, Brazil, and a severe pruning system, the cv. ‘Roxo de Valinhos’
develops one fruit per node, reaching an annual shoot growth of 189 cm (Alvarenga-
Gonçalves et al., 2006). In Japan, in the cv. ‘Masui Dauphine’ under greenhouse
cultivation, one fruit is borne on every node of the shoots, and the harvesting period
is extended from the base to approximately the 20th node (August to November in
the northern hemisphere) (Hosomi et al., 2013). Under Mediterranean climate con-
ditions and light pruning, the percentage of nodes with fruit in the Main crop of five
fig varieties ranged between 50.2 and 88.5% (Gaaliche et al., 2011), whereas in the
temperate climate conditions of central Argentina and intense pruning, the percent-
age of nodes with fruit ranged between 45.0 and 49.9% (Micheloud et al., 2018).
Concerning harvest distribution, in this region of Argentina and under severe prun-
ing, it is abundant from January to the end of March (Micheloud et al., 2018) and
declines during April and May (Fig. 8.4).
In the fig, the annual duration of the harvest period ranges from 8 to 21 weeks,
with modification of its ending but not of its beginning date when different years are
compared. Furthermore, the duration of harvesting shows a great linear relationship
with the annual fruit yield, which, in the central area of Santa Fe, Argentina, ranges
between 4.43 and 12.1 t ha−1 (Micheloud et al., 2018). This fruit yield is in accor-
dance with that observed in different areas of Brazil using the cv. ‘Roxo de Valinhos’
and severe winter pruning (Leonel & Tecchio, 2010; Nienow et al., 2006).
The duration of the harvesting period and fruit yield are also affected by tree age.
In Santa Fe, the longest harvesting period was observed in the fourth growing sea-
son after planting, with a later decline (Micheloud et al., 2018). This effect was not
observed under tropical conditions of Minas Gerais, Brazil, where fruit production
and the duration of the harvesting period are stabilized after the fifth year of planta-
tion (Alvarenga-Gonçalves et al., 2006). The duration of the harvesting period is
also affected by meteorological conditions, decreasing one week for every 5–6 rainy
204 N. Micheloud et al.

3.0

Brown Turkey
2.5 Guarinta

2.0
Kg.plant-1

1.5

1.0

0.5

0.0
Dec Jan Feb Mar Apr May

Fig. 8.4 Monthly distribution of the annual fruit harvest (Kg plant−1) of two fig (Ficus carica L.)
cultivars, ‘Brown Turkey’ and ‘Guarinta’, under a severe pruning system with annual renewal of
the canopy in the central-east area of Santa Fe, Argentina. The vertical lines indicate the standard
error. Data are the mean values of ten years of experimentation. (Adapted from Micheloud
et al., 2018)

days (>75 mm of accumulated precipitations) from January to May (Micheloud

et al., 2018).
It is possible that, under severe pruning, the development of the root system
makes the fig tree too vigorous, with a later fruit set that is unable to reach maturity
under temperate climate conditions. Thus, reducing the harvest period occurs not
because of the absence of fruits but because of the lack of ripeness (Micheloud
et al., 2018). In contrast, the ripening of later fruits normally occurs under tropical
climate conditions (Alvarenga-Gonçalves et al., 2006), explaining the different evo-
lution of fruit yield and the duration of the harvesting period with tree age under
severe pruning intensity under temperate and tropical climates. Furthermore, the fig
fruit has the particularity that, to ripen successfully, it needs high temperatures,
preferably above 30 °C, which explains why it is common to observe a large number
of immature fruits during the autumn period in temperate zones (Lavín & Reyes,
2004). It also explains why the harvest period ends despite fruits on the plant.

8 Fig Production Strategies

Fig trees can be adapted to marginal conditions and tolerant of high soil calcium
content, salinity, and drought (Golombek & Lüdders, 1990). As a result, figs have
recently attracted a great deal of attention and are becoming widespread worldwide.
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 205

In Asia, several countries such as China, Japan, Thailand, Indonesia, and Malaysia
have started to plant figs (Isa et al., 2020). In Mexico, fig cultivation has also
increased both under open field and greenhouse conditions because it responds
favorably to agronomic management (Mendoza-Castillo et al., 2017) and provides
economic viability to the rural sector (Macías et al., 2014). However, since open
field production in Mexico has a higher incidence of pests and diseases, fruit rotting
during the rainy season, and frost damage, in comparison with production under
greenhouse conditions, the latter is becoming more expanded than the former
(Mendoza-Castillo et al., 2017).
Currently, contrasting strategies can achieve fig production (Table 8.1). One of
these strategies is the traditional system of semi-arid regions, mainly used in
Mediterranean countries, characterized by open field cultivation, rainfed conditions,
and low tree density (156 pl ha−1). Other features of this system are a slight pruning,
one (Main) or two (Breba and Main) crops per year according to the variety, low
shoot growth around the year (6.8–to 14.2 cm), and several shoots per plant (non-
controlled), three (Breba crop) to six (Main crop) fruits per shoot, and a fruit yield
of 7–8 t ha−1 for the Breba crop (Gaaliche et al., 2011) and 15 t ha−1 for the Main
crop (Melgarejo-Moreno, 1999). In addition, the use of different organic and inor-
ganic mulches under rainfed conditions improves soil moisture content, the number,
and width of leaves, shoot length and diameter, and fruit quality, and results in
milder and more stabilized soil temperature throughout the year (Jafari et al., 2012).
Figs can also be cultivated under an intensive system in greenhouse conditions,
where plants are spaced at 1.6 × 0.5 m (12,500 plants ha−1) using 40-L containers
per plant, substrate (soilless cultivation), and conducted with 3–8 stems per plant (6
shoots on average with 20 fruits per shoot), with severe pruning and annual renewal
of the canopy. In this strategy, the nutritive solution is applied by drip irrigation dur-
ing the whole growing season. In Chapingo, Mexico, by using the native cultivar
‘Netzahualcoyotl’, the potential yield of this production system is over 60 t ha−1.
This fresh fruit yield results up to 20 times higher than that achieved by open field
plantations in the same area of Mexico. Total dry mass production reaches near
30 t ha−1, and dry matter partition to fruit ranges from 38 to 57% between extreme
treatments (3 and 7 shoots per plant, respectively). In the northern hemisphere, the
harvesting period lasts 3 months, from July to September (Mendoza-Castillo
et al., 2017).
In Japan, fig cultivation under greenhouse is widespread using the “straight line
training” system, adopted mainly for the cv. ‘Masui Dauphine’. The system involves
bilateral horizontal limbs of 40 cm height with shoots growing upward from them.
Shoots are cut off at the end of the growing season, similar to that performed in
grapevines. This system greatly improves cultural practices such as pruning, dis-
budding, and harvesting (Hosomi et al., 2013).
Finally, in the central area of Santa Fe and other regions of Argentina, growers
adopt an intermediate fig production system (Table 8.1). This system is character-
ized by open field conditions with complementary drip irrigation, a tree density near
206 N. Micheloud et al.

Table 8.1 Description of different production systems for the fig tree crop
Traditional Intermediate Intensive
Location M’hamdia (Tunisia) Santa Fe (Argentina) Chapingo (Mexico)
Coordinates 35°55′N, 10°34′E, 18 31°26′ S, 60°56′ W, 20 19°20′N, 98°53′W,
masl masl 2240 masl
Climate Semi-arid. Mild winter Temperate humid with no –
and hot, dry summer. dry season
Annual 439 1100 –
precipitation
(mm)
Soil Silty-clay, saline Sand-loamy Soilless (40 l plant−1
container)
Cultivation Open field Open field Greenhouse
methods
Irrigation Rainfed Complementary drip Drip irrigation with
irrigation nutrient solution
Cultivar ‘Bither Abiadh’, ‘Brown Turkey’, ‘Netzahualcoyotl’
‘Besbessi’, ‘Khedhri’, ‘Guarinta’
‘Chetoui’, ‘Zidi’
Crops per year 2 (Breba and Main 1 (Main crop) 1 (Main crop)
crops) (San Pedro cvs.)
1 (Main crop) (Smyrna
cvs.).
Plant density 8.0 × 8.0 m 4.0 × 2.5 m 1.6 × 0.5 m
(156 pl ha−1) (1000 pl ha−1) (12,500 pl ha−1)
Plant age 15 7 –
Shoots per plant Non-controlled 25–30 3–8
Shoot length 6.8–14.2 160 –
(cm)
Fruit per stem 3 (Breba crop) 25 20
6 (Main crop)
Pruning system Light Annual renewal of the Annual renewal of the
canopy canopy
Dry matter – – 15.25–29.6
production
(t ha−1)
Dry matter – – 38–57%
partition to fruits
(%)
Fruit production 7–8 (Breba crop) 7–15 40.5–109.5 (60 on
(t ha−1) 15 (Main crop) average)
Fruit harvest 2 (June-July; Breba; 4–5 (January-May; south 3 (July-September;
period (months) north hemisphere) hemisphere). north hemisphere)
3 (July-September; Induction of last fruit
Main crop; north maturation during autumn
hemisphere) (May).
Fresh fruit weight – – 52.37–58.45
(g fruit−1)
Fruit length (cm) – – 6.57–7.56
Traditional: Adapted from Gaaliche et al. (2011); Intermediate: adapted from Micheloud et al.
(2018); Intensive: Adapted from Mendoza-Castillo et al. (2017)
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 207

Fig. 8.5 Fig crop for fresh

market and industry after
winter pruning with an
annual canopy renewal in
central Argentina. Tree
density 6 × 3 m. (Photo:
Obrador, C.)

1000 pl ha−1 (4.0 × 2.0 m; 4.0 × 2.5 m), severe pruning with annual renewal of the
canopy, 25–30 shoots per plant, 25 fruits per shoot, and a fruit yield of 7–15 t ha−1.
In addition, the harvesting period is extended from December to May (south hemi-
sphere) (Micheloud et al., 2018). However, the fig tree responds favorably to an
increase in nitrogen fertilization, with farms that achieve commercial yields up to
35 t ha−1, even with a lower tree density (6 × 3 m, 555 pl ha−1) (Fig. 8.5).
Unlike many other fruit trees (Berman & Dejong, 2003; Bussi et al., 2005;
Mahmood et al., 2000), the fig tree has the particularity that vegetative growth does
not affect the fruit set of the Main crop (Hosomi et al., 2015). Consequently, the fig
tree has great reproductive stability under different growth conditions, and, accord-
ing to our observations, even an increase in the tree vigor does not delay the begin-
ning of harvesting.
208 N. Micheloud et al.

8.1 Stimulation of Fruit Growth and Ripening

Since the third Century BC, it has been known that fig growth and ripening could be
stimulated by applying olive oil and other vegetable oils to the ostiole of the fruit
(oleification) (Chessa, 1997; Hirai et al., 1966) because this induces ethylene pro-
duction. Furthermore, fruit growth and earlier ripening can also be stimulated by
auxin (Miller et al., 1987) and ethephon. However, it has been recently demon-
strated that ethylene production in response to these compounds is induced by
1-methyl cyclopropane and inhibited by propylene, indicating a negative feedback
regulation mechanism (Owino et al., 2006).
These techniques used for the anticipation of fig fruit ripening have been used for
the Breba crop because the advance of its maturity is justified by the higher prices
of the earliest fruit in the fresh market; furthermore, the uniformity of maturity
reduces the labor costs of harvesting (Melgarejo-Moreno, 1999).
It is essential to consider that, in the temperate climate conditions of Santa Fe,
Argentina, near eight fruits per shoot are not able to reach maturity because they set
later in the growing season (Micheloud et al., 2018), and the fig requires lots of heat
units to reach maturity and achieve good fruit quality (Stover et al., 2007), limiting
fig cultivation in low-temperature regions. Furthermore, these late fruits that remain
on the plant are later affected by frost and then abort, resulting in a considerable loss
of fruit yield because they represent up to one-third of the total fruit sets annually by
the fig tree (Main crop).
The stimulation of fruit growth and ripening used traditionally for the Breba crop
has been evaluated with satisfactory results in a temperate climate to ripen the fruits
that set later and remain green at the end of the growing season, increasing fruit
yields under a severe pruning system. Under unfavorable autumn temperature con-
ditions, olive oil and ethephon (200–400 ppm) can ripen the fruits, allowing them to
reach similar size and weight but lower total soluble solid content (°Brix) than fruits
that ripen early during normal temperature conditions (unpublished data). Ethephon
is applied to the plant but directed to the most basal fruits of each current shoot. A
weekly application is carried out for 3 weeks, and one-third of the remaining fruits
are treated at each application. Treated fruits ripen during the week of treatment,
previous to the next ethephon application (unpublished data).

9 Conclusions

In temperate humid areas, such as Santa Fe, in central Argentina, fig tree cultivation
for the fresh market can be carried out successfully. The main strategies that have
been proved to be more effective in obtaining high-quality fruit and yield are; (i)
high-density plantations (>1000 pl ha−1); (ii) severe pruning with annual canopy
renewal; (iii) stem growth stimulation through irrigation and fertilization, which
allows a long harvest period of the Main crop (3–5 months); (iv) fruit ripening
8 Agronomic Strategies for Fig Cultivation in a Temperate-Humid Climate Zone 209

stimulation by fruit-localized ethephon application either to advance fruit harvest

prior to a rainy period or for the ripening of the fruits that remain green on the plant
at the end of the growing season, and (v) high-technology application such as green-
house cultivation with plantations over 10,000 pl ha−1 and fruit yield that could
exceed 100 t ha−1. However, further research on fig cultivation in temperate-humid
areas should be carried out to adjust these incipient production strategies.

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Chapter 9
Cultivars and Agriculture Practice of Fig
(Ficus carica)

Walid Nosir

1 Introduction

The fig tree has a lengthy history; archaeologists have discovered evidence of its
cultivation dating back to 5000 BC. Fig trees are tiny, warm-climate trees that could
grow practically everywhere, with some types surviving in temperatures (−12 to
−6 °C.). For around 15 years, fig-trees will yield well. The fig is mentioned in the
holy books, such as Bible and the Quran. It is also shown in Egyptian hieroglyphics.
It is adaptable to moderate climates and has been highly valued and widely culti-
vated in most Mediterranean countries for millennia. The first figs were reportedly
delivered to Parris Island, South Carolina, in 1577, where they were found thriving.
They were discovered 2 years later near Saint Augustine, Florida. Figs were intro-
duced to Virginia from the Bermudas in 1621 and were noted to be nutritious there
in 1629. In 1560.
There are four types of figs in the horticultural world. Capri figs yield a small,
non-edible fruit, but their blossoms produce pollen. Pollen from Smyrna and San
Pedro figs is necessary to fertilize their flowers (Ferguson et al., 1990; McEachern,
1996). Smyrna figs (Calimyrna figs in the United States) produce huge, edible fruit
with viable seeds (Condit, 1955). Normal fruit development necessitates pollina-
tion. San Pedro figs have two crops: a parthenocarpic breba crop (first crop) and a
pollinated main crop (Tous & Ferguson, 1996). Although several common figs gen-
erate a breba crop, one trait that distinguishes San Pedro-type figs is the setting of a
breba crop (Flaishman et al., 2008). The common fig, which includes the varieties
‘Bourjasotte Noire,’ ‘Col de Dame Noire,’ and ‘Noire de Caromb,’ develops parthe-
nocarpically and rarely requires pollination (Ozeker & Isfendiyaroglu, 1997). As a

W. Nosir (*)
Vertical Farm Academy, Newburgh, IN, USA
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 215
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_9
216 W. Nosir

result, the fruit usually has underdeveloped, sterile pips (Caliskan & Polat, 2008).
Some common fig cultivars, such as Bourjasotte Noire, shed all or virtually all of
the breba crop‘s fruit buds before producing a main productive crop. ‘Noire de
Caromb,’ show almost complete parthenocarpic growth in both crops (Ferrara &
Papa, 2001). Fig trees can bear two crops/ a year (Bostan et al., 1997). The first crop,
the breba, is borne laterally on the previous season’s growth from buds produced in
the leaf axils. The buds start developing with the onset of the following spring, and
fruit usually matures within the first month of summer. The second crop, consider
the main crop, also arises from buds in the axils of leaves but on shoots of the cur-
rent season. Fruit maturation may start in the middle of summer and can last several
weeks (Flaishman et al., 2008). Production practices must be created to produce figs
profitably for the local growth circumstances. Understanding the phenology of
‘Bourjasotte Noire,’ ‘Col de Damme Noire,’ and ‘Noire de Caromb’ as expressed
under local growing conditions is required for tree training using pruning and rest
breaking agents (RBAs). When left to grow organically, fig tree canopies can reach
a height of 15 meters. Allowing trees to reach this size is ideal for producing dry figs
harvested off the ground. However, in most modern orchards where figs are grown
for fresh consumption, trees are pruned to keep their height under three meters to
provide fruit pickers easy access (Flaishman et al., 2008). Fig tree leaves have been
shown in several experiments to lower blood sugar and total cholesterol while
increasing antioxidant levels.
According to FAO, the area planted with F. carica in 2021 exceeded 457,737
hectares, with an output of over one million metric tons per year (FAOSTAT).
Table 9.1 presents the top fig-producing countries. The countries of the European
Union produce the majority of the world’s F. carica. Fig fruits are essential compo-
nents of the Mediterranean diet, which is often regarded as one of the healthiest on
the planet. It is a dietary habit that’s also linked to longevity. The cultivation of fig
trees is widespread. They thrive in the Mediterranean and are well-adapted to
drought and high temperatures. The nations with the most increased production of
F. carica are Turkey and Egypt, Algeria, Morocco, and Iran, which account for 43%

Table 9.1 Top fig growing countries

Annual production of fresh figs in
Rank Country metric tons
1 Turkey 262,644
2 Egypt 203,238
3 Morocco 76,625
4 Iran 75,834
5 Algeria 72,898
6 Greece 50,368
7 Syria 44,032
8 United States 43,001
9 Spain 42,856
10 Tunisia 25,786
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 217

of global production. Turkey is the most important country. Austria, Spain, Italy,
and the Netherlands are the biggest exporters of F. carica, whereas the Netherlands
is the primary importer. France, Germany, Austria, and the United Kingdom are
home to F. carica. Commercial fig production is carried out in Gozlekci, Gok, and
Yilmaz. California, Australia, and South America, among other places, have a
Mediterranean climate (Pereira et al., 2020). Figs have recently gained much popu-
larity and are grown worldwide. Several Asian countries, including China, Japan,
and Thailand, have begun cultivating F. carica.
The commercial was assessed by Lianju, Weibin, Kai, Zhifeng, and Yelin. The
output of figs grew in China due to scientific studies into this fruit, including cultivar
selection, cultivation methodologies, training, pruning techniques, and soil manage-
ment. In addition, the production of figs was aided by fertilizer, irrigation, process-
ing techniques, and medical applications. As a result, the fig is widely favored by
growers. Furthermore, before 1985, the area dedicated to fig cultivation was approx-
imately 350 hm2, and currently, there are 6667 hm2 available, which is 20 times
more than 20 years ago.
Meanwhile, the figs industry is growing. However, the industry is only concen-
trated in a few areas in Japan, with “Masui Dauphine” being the dominant cultivar.
Thailand’s fig industry concentrates on developing new traditional agricultural
goods to sell in local markets.
The Ficus carica L. tree is an underutilized and minor fruit species in Malaysia,
with cultivation primarily for hobby purposes. Currently, there are only a few F. car-
ica orchards scattered over Malaysia. However, although F. carica farming is still in
its infancy, it is progressively gaining traction among those captivated by this
ancient plant (Vemmos et al., 2013). As a result, F. carica has been proposed as an
alternative high-value crop that provides farmers with a new source of income
because it is easier to produce, could be harvested in a short period, and provides a
faster return on investment than fruit crops. Furthermore, the cultivation of F. carica
has grown worldwide, owing to its uses as a food, medicine, and cultural symbol.
This issue has sparked much interest in looking into its production processes.

2 Fig Cultivars

The mandatory outcrossing in this species increases genetic variety in figs, resulting
in developing new varieties with potentially advantageous features from seeds.
Because fig is easily propagated through cuttings and is routinely re-propagated to
preserve suitable cultivars, natural mutations within a cultivar could produce signifi-
cant phenotypic heterogeneity. The practice of naming excellent fig cultivars dates
back to the fourth century BCE. Flaishman et al. (2008) listed 607 varieties that
produce fruit. However, only a few cultivars are used in commercial production.
The California fig industry, for example, is based mainly on five cultivars:
218 W. Nosir

‘Calimyrna’ (‘Sarilop’), ‘Adriatic,’ ‘Mission,’ ‘Brown Turkey,’ and ‘Kadota’

(California Fig Advisory Board, 2006; California Fresh Fig Growers
Association, 2006).
Leaf morphology, plant vigor, external and internal color, flavor, percentage sol-
uble solids, titratable acidity, seed properties, fruit shape, skin thickness, ostiole
diameter, and fruit production time are all qualities cultivars differ. Traditionally,
distinct cultivars were classified based on fruit and tree features. This method is
practical and sensible, especially when marketing fruit or selecting planting mate-
rial. However, other cultivars have similar descriptions, and a single genotype is
thought to be associated with dozens of names in certain circumstances (Flaishman
et al., 2008). More precise data on brebas, main-crop fig leaves, and fruit has been
proven to characterize practically all cultivars in the Extremadura collection in
Spain (Giraldo et al., 2010). This description has the advantage of requiring less
specialist equipment, but it is more prone than molecular markers to be altered by
environmental variability. Isozyme analysis has been used to sort genotypes for
many years among molecular approaches. Even though isozymes have been used to
differentiate fig cultivars in several studies, only a few cultivars have been studied in
practice. Pereira et al. (2020) reported six enzyme systems to identify a more sig-
nificant number of cultivars, but just the three most helpful enzyme systems were
needed to generate distinct patterns for each of the 31 genotypes studied. Using 25
primers to synthesize random amplified polymorphic DNA, these researchers dis-
covered comparable discriminating (RAPDs). DNA microsatellites, also known as
simple sequence repeats (SSRs), have proven to be particularly valuable in finger-
printing genotypes in various animals. Harrison et al. (2003) studied RAPDs, inter
simple sequence repeats (ISSR), and microsatellite markers for molecular charac-
terization of 30 cultivars, finding that RAPDs were the least effective technique for
distinguishing fig genotypes. In a study of 70 fig accessions at the Conservatoire
Botanique National Méditerranéen de Porquerolles in France (Harrison et al., 2003),
52 genotypes were discovered using five SSR loci. While the authors speculated
that these were most likely 52 different genotypes with several duplicates labeled
differently, they also proposed that more markers be used. Six SSR loci were used
to analyze 75 fig accessions, resulting in the identification of 72 genotypes (Khadari
et al. 2005). Giraldo et al. (2010) used 25 polymorphic SSR loci to describe 15
accessions. The SSRs revealed an average of 3 alleles per locus and 11 distinct
genotypes among the 15 accessions, indicating a small genetic base in cultivated
figs. 16 SSR loci were recently utilized to analyze 181 fig accessions in the
Germplasm Repository in Davis, California, yielding 128 distinct genotypes
(Aradhya and Stover, personal comm.). In an ideal scenario, fig researchers world-
wide would agree on a set of markers that could be used to verify the authenticity of
figs across different collections.
Characteristics that should be considered in the selection of fig cultivars include
the following: cold hardiness, the ability to set fruit without pollination (partheno-
carpy), the fruit having a closed eye, or ostiole, a long peduncle that allows the fruit
to droop and shed moisture, and a green skin on the fruit to minimize bird herbivory.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 219

2.1 Fig Cultivars

Alma This cultivar is reported to produce high yields. Alma is moderately cold
tolerant and produces a medium-sized fruit with brown skin and a light tan pulp
(Fig. 9.1). The eye is medium open, and the fruit has few seeds and is very sweet.
Fruit ripens from late July through August and is good fresh or processed. Considered
a high-quality fig tree, the Alma was bred at Texas A&M University and has adapted
well to the southeast. Alma trees have dark green foliage and require full to partial
sun. Trees thrive in well-drained, sandy, moist, loam soils but are highly tolerant of
poor soil conditions. The Alma is an extremely productive tree and produces figs
that ripen in late June and again in August. The fruit is small to medium and has a
pear shape, golden-brown skin, and amber pulp. Alma figs are sweet, delicate
caramel-flavored figs best served fresh, dried, and preserved.
A small to medium, golden-brown, pear-shaped (pyriform) fig with amber/light yel-
low pulp. Sweet and delicate flavor. Well-adapted in the Southeast. Highly resistant
to fruit rots. Wood is very hardy. Excellent quality, succulent and sweet, with barely
noticeable seeds and a well-sealed eye. Very productive, compact tree is cold, hardy,
and has a prolonged spring dormancy.
Black Spanish (California Brown Turkey, San Pedro): This cultivar yields
medium-sized, purplish-brown fruit with a red center (Fig. 9.2). Black Spanish has
an eye open. Fruit ripens in June and in August.

Brown Turkey (Brunswick, Eastern Brown Turkey, Harrison, Lees Perpetual,

Ramsey, Texas Everbearing). This cultivar is probably the most popular fig in the
southeastern United States. In the spring, a small fruit crop (called breba fruit) rip-
ens in July, followed by the main crop one month later (Fig. 9.3). The fruit is

Fig. 9.1 Fig flavors, pulp and skin color examples. (Credit: https://mountainfigs.net/)
220 W. Nosir

Fig. 9.2 Fig Alma

Fig. 9.3 Black Spanish Fig

medium-large in size and has bronze skin and amber pulp. The fruit has a small to
the medium closed eye and is good fresh, or processed.

Celeste (Blue Celeste, Celestial, and Little Brown Sugar). This cultivar is probably
the second most common fig in the southeastern United States. Celeste is fairly cold
and hardy; the fruit is small to medium in size and purplish bronze to light brown
(Fig. 9.4). Celeste has a closed eye and begins ripening in early July.

Champagne (Golden Celeste). This newly released cultivar from Louisiana State
University produces a medium-sized fruit with yellow skin, tan-colored pulp, and a
closed ostiole. Fruit ripening is in early July (Fig. 9.5).
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 221

Conadria (Genoa). This cultivar is a vigorous tree that produces green to yellow
fruit (Fig. 9.6). The flesh of the fruit is pink to red with a good flavor. The eye is
small and tight. Fruit ripening is in June and again in August. Conadria fruit is good
fresh and excellent dried.

Green Ischia (Ischia Green, Ischia Verte, and White Ischia). This cultivar produces
a green small to medium-sized fruit with a strawberry center and a closed eye
(Fig. 9.7). Fruit of Green Ischia ripens in late July to early August.

Hunt This cultivar is very cold and hardy and produces a small pear-shaped, violet-
brown fruit with a long neck (Fig. 9.8). Hunt has a closed eye, amber-colored flesh,
and few seeds; the fruit ripens in July.

Jelly (Mary Lane Seedless). This cultivar produces a longnecked, yellow fig
medium in size with clear amber flesh and very few seeds (Fig. 9.9). Fruit of Jelly
is good for eating fresh and preserving, although the skin is soft. Jelly ripens from
late July to August.

Kadota (Florentine). This cultivar produces a medium-large, yellow fruit with an

open ostiole partially sealed with a honey-like substance (Fig. 9.10). Fruit quality
declines with extremely wet weather. Although Kadota figs can be eaten fresh, they
are better suited for canning and preserves. Fruit ripens in July.

LSU Gold This cultivar is a Louisiana State University release producing a large,
yellow fig with pink-to-red pulp. The fruit of LSU Gold should be picked as soon as
it is mature because this fruit has an open eye, and fruit spoilage may occur
(Fig. 9.11). Fruit ripens in July through August. Fruit is of good quality for eating
fresh and for preserving.

LSU Purple This cultivar is a Louisiana State University release that produces a
medium-small, glossy purple fig with amber-to-pink flesh and a closed eye

Fig. 9.4 Celeste Fig

222 W. Nosir

Fig. 9.5 Champagne

(Golden Celeste) Fig

Fig. 9.6 Conadria

(Genoa) Fig

(Fig. 9.12). The main crop ripens in August, but some fruit can ripen well into the
fall. Fruit is of good quality for eating fresh and for preserving.

Magnolia (Brunswick, Madonna). This cultivar is common in some southeastern

United States but not in Florida. Magnolia is cold, hardy down to 50 °F, and often
produces the largest fig. The fruit is asymmetric, bronze in color, and has an open
ostiole (Fig. 9.13). The flesh is amber to strawberry in color. Ripening is from mid-
July through August. Fruit should be picked early because it may split and turn sour
under wet conditions.

Mission (Black Mission, Franciscana). A large, black fig with reddish-pink pulp
(Fig. 9.14). The mission is an ever-bearing fig that produces fruit from summer to
winter. Not sufficiently cold hardy for the southeastern United States.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 223

Fig. 9.7 Green Ischia Fig

Fig. 9.8 Hunt Fig

O’Rourke (Improved Celeste). This cultivar is a Louisiana State University fig

that produces a small to medium-sized fruit brown with tan pulp (Fig. 9.15). The eye
is partially closed with the aid of a honey-like substance. Fruit ripens in early July.
Fruit is of good quality for eating fresh and for preserving.

Osborn Prolific (Arachipel, Hardy Prolific, Neveralla, Osborne, Rust). This culti-
var produces a medium-large fruit with reddish-brown skin and light-colored flesh
(Fig. 9.16). The fruit is sweet with few seeds. The eye is partially closed. Osborn
Prolific is reported to perform better in cooler climates. Fruit ripens in August, and
the fruit is best eaten fresh.
224 W. Nosir

Fig. 9.9 Jilly Fig

Fig. 9.10 Kadota Fig

Pasquale (Natalino, Vernino). This cultivar produces a sweet, small, purple fig
with amber-to-pink pulp (Fig. 9.17). Pasquale ripens from late November to
December and is often damaged by frost. However, Pasquale is not cold-hardy. A
medium-large fig that is greenish-yellow in color with light strawberry pulp. The
fruit has a closed ostiole. Tena thrives in hot, dry weather. The fruit is good for eat-
ing fresh or preserving.

Tiger (Giant Celeste). This new cultivar released by Louisiana State University has
large brown fruit, yellow pulp, and a partially closed eye (Fig. 9.18). Fruit of Tiger
ripens in early July.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 225

Fig. 9.11 LSU Gold Fig

Fig. 9.12 LSU Purple Fig

3 Production Process

F. carica cultivation is rapidly increasing in the open field and under greenhouse
conditions because it is a species that responds well to agronomic management,
making it an excellent production alternative. However, F. carica planted in the
open field often encounters high pest and disease incidence, rotting of the fruit dur-
ing the rainy season, and poor plantation management. These harmful conditions
226 W. Nosir

Fig. 9.13 Magnolia fig

Fig. 9.14 Mission Fig

will have a significant impact on crop growth and fruiting. Thus, the shielded cul-
ture of F. carica will be the solution amid severe growing conditions. Greenhouse
horticulture is the most intensive mode of crop production, yielding up to 10 times
more per cultivated unit area than a field crop. Growing crops in a protected envi-
ronment protect them from natural hazards. It allows for artificial manipulation of
the crop micro-environment to facilitate optimal plant performance, extend produc-
tion duration, induce early flowering, and improve production and product quality
(Figs. 9.19 and 9.20). Protected cultivation allows for year-round production, which
is impossible in open field farming due to excessive rainfall and wind, especially in
tropical locations (Pereira et al., 2015).
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 227

3.1 Nutrient Control

Fertilization management is critical in agriculture because of economic and envi-

ronmental concerns. Over the last few decades, agricultural crop nutritional require-
ments mismanagement has resulted in significant health and environmental
concerns. The fertilizer should be applied based on the location’s variety, soil type,
and climate. The nutritional needs of the plants are usually met by proper fertiliza-
tion. However, it is vital to apply fertilizers sparingly to lessen fertilization’s envi-
ronmental impact. In a previous study by Lianju et al., nitrogen fertilizers such as
urea and NH4HCO3 were utilized in orchards where the soils were infertile. The
weather was less rainy during the early growing season. Because fig fruits at the
lower regions of shoots began to mature while those at the top were just at the stage
of differentiation, potassium and phosphate fertilizers were frequently sprayed
more in fig orchards and sooner than in other fruits. One month before harvesting
summer and fall fruits, potassium and phosphate fertilizers were often applied.
According to a study on the effects of potassium fertigation on yield, marketable
output, and fruit size in two different fig cultivars, ‘Black Mission’ and ‘Sierra,’
considerable fertigation of potassium may enhance the marketable yield; of ‘Black
Mission.’ And ‘Sierra’ fresh fig cultivars are critical for new fig growers because the
commercial output is revenue (Islam et al., 1997). The fertigation treatment ‘Sierra’
figs also boosted fruit size, and this rise is linked to high leaf K concentrations. The
administration of correct fertilizers could result in increased growth with enhanced
fruit output and quality because any nutrient, whether lacking or excessive, could
result in a drop in crop yield and inferior fruit quality.
Little research has been done on the use of fertilizers for figs. However, there is
certain basic information that is useful. Fig trees characteristically produce good
crops under relatively low fertility levels. However, increased vigor and fruiting can
often be achieved by fertilizers. This is particularly true where figs are grown on
infertile, sandy soils and grown in competition with lawn grasses and ornamental

Fig. 9.15 O’Rourke

(improved Celeste)
228 W. Nosir

Fig. 9.16 Osborn

Prolific Fig

Fig. 9.17 Pasquale Fig

plants. Fig trees habitually grow vigorously, and the leaves are large. Below-normal
leaf size and terminal growth indicate the need for fertilizing. Fig trees should be
fertilized, as should other fruit trees such as peaches and plums, except that fig trees
rarely experience iron and zinc deficiencies. Nitrogen is usually the only mineral in
short supply. For large fig bushes, 1 to 1.5 pounds of nitrogen per tree per year was
applied in late 18 AGR. Winter or early spring is generally adequate, but more fre-
quent applications may be required in heavy rainfall areas. The fertilizer amount can
be increased or decreased as indicated by the bush size and the response obtained.
In those areas where complete fertilizers are needed, a mixed fertilizer applied in an
amount that will contain 1–2 pounds of nitrogen should be used (Fig. 9.21). The
phosphorus and potassium ratios to nitrogen should be about the same as commer-
cial acreages.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 229

Fig. 9.18 Tiger Fig

Fig. 9.19 Fig propagation steps using one-year cuttings

3.2 Water Requirement

Because water is a scarce resource in many parts of the world, producing more with
less water in agriculture is critical. Irrigation water is constrained by two factors:
water scarcity and inefficient water utilization. The irrigation system is one of the
most critical factors determining the yield and quality of agricultural produce in a
greenhouse farming system. Water should be given in the right amount and at the
right time. As a result, water management is critical for avoiding plant moisture
230 W. Nosir

Fig. 9.20 Fig air layering

stress during the growth stages of the crop (Saddoud et al., 2008). As a result, the
development of water-efficient agriculture and crop yield and water usage efficiency
improvements have much potential as practical approaches for developing sustain-
able agriculture in irrigated areas.
F. carica plants prefer drier growth conditions than most fruit trees. However,
this plant still needs adequate water for optimal performance and production. The
frequency is determined by the trees’ size, vigor, soil type, and rainfall. Because
F. carica trees are susceptible to root rot, it is best to prevent severe moisture swings,
such as excessive irrigation water, as F. carica cannot thrive in these conditions.
Caruso et al. (2022) collaborated on a study to assess the gas exchange and growth
response of F. carica plants to water deficit and relief. According to the study, mild
water deficiency substantially impacts shoot growth, leaf biomass, and gas exchange
parameters. El-Shazly et al. (2014) got similar results on many F. carica cultivars
grown under various irrigation regimens. Compared to 100% ETc, watering at 75%
ETc and 50% ETc resulted in lower growth rate values.
Treating the crop with water without first understanding the parameters involved
will result in drainage and salinity issues and a failure to reap the benefits of irriga-
tion. As a result, knowledge of proper irrigation management is critical for the
growth of F. carica, as it will result in increased crop production and more efficient
use of water resources.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 231

3.3 Soil Requirement

According to Aljane and Ferchichi (2006), the fig could be cultivated on various
soils, including light sand, rich loams, clay, or limestone, as long as there is suffi-
cient depth and proper drainage. Excessive plant growth is encouraged at the
expense of fruit yield in heavy, damp soil. Soils with a high acidity level are inap-
propriate. Between 6.0 and 6.5 is the optimal pH range. Figs thrive in soils that are
well-drained, fertile, and high in organic matter (Bostan et al., 1997). They indicate
that fig trees may adapt to various soils, but the best are those with a clayey-sandy
texture, high organic matter, and a pH of 6.0–6.8. In poorly drained soils, root rots
can occur, and in overly dry soils, plants go into a dormant state, producing few
leaves and no fruits. In recent years, the harvested area and volume of fig production
have remained stable globally, with Turkey, Egypt, Morocco, Algeria, and Iran serv-
ing as the primary producers. Fig cultivation has increased tendency in some nations,
such as Mexico, with an average yearly growth rate of 6% in harvested area and
13% in output volume.
In 2017, 1440 ha of land were harvested in Mexico, with an average yield of 5.6 t
Darjazi, 2011. Under greenhouse conditions, however, products of above 100 t ha1
have been reported. Crop development can be influenced by various physical, chem-
ical, and biological factors, including plant nutrition. In plants, nutrients play a cru-
cial and unique role. When one of these elements is not present in sufficient amounts
in the plant’s tissues, it causes alterations in the plant’s metabolism, affecting veg-
etative growth.

Fig. 9.21 Fig growing in containers using soilless culture technique

232 W. Nosir

4 Fig Production in Soilless Culture

4.1 Propagation

Starting a fig tree from fig cuttings is easy and can be done in three ways. Each of
these ways of rooting figs is basic and straightforward, and the weather conditions
in your area will likely determine your choice during the dormant season. Fig prop-
agation layering, the first method of propagating fig trees outside, relies on tempera-
tures that never drop below freezing during the dormant season. Ground layering is
a method of rooting figs that involves burying a piece of a low-growing branch
15–20 cm of the tip visible above ground, allowing the buried portion to root, and
then removing the buried portion from the parent tree. This is the basic way of fig
growth; keeping the ground clean while the branches root can be challenging.
Finally, planting fig cuttings in the garden fig cuttings is a more popular method of
rooting figs outside. Take fig cuttings from little branches that are 2–3 years old, late
in the dormant season, when the threat of frost has passed. They should be 12–34
inches thick (1.3–1.9 cm) in diameter, about the breadth of your pinky, and 8–12
inches (20–30 cm). The bottom end should be flat, while the branch tip should be
one-sided. Use a sealant on the slanted end and rooting hormone on the flat end to
prevent disease. Fig cuttings can grow 36–48 inches in a year (91–122 cm.). The
new trees will be ready to be transplanted during the following dormant season.

4.2 Soilless Media for Growing Fig Indoor

The variety of figs, the pH of irrigation water, the cost, the product’s shelf life, the
type of system utilized, and the grower’s preference go into selecting a growing
medium. When selecting a medium, a grower should search for specific traits. For
example, soilless media, like soil, must offer oxygen, water, nutrients, and support
for plant roots (Stone, 2014).
All or some of the following properties should be present in an optimal grow-
ing medium:
aeration and drainage are also important. While the medium must be able to
retain water, it must also be able to drain well. Excessively fine materials should be
avoided to minimize water retention and a lack of aeration within the medium. The
soilless media should contain the following characteristics;
–– Durability. The medium must be long-lasting. Avoid using soft aggregates that
readily dissolve.
–– Porosity. Between cycles, the medium must remain wet from the nutrient flow
long enough for the plants to receive their essential nutrients.
–– Sterile. Diseases and pests will be less likely to spread if the growing medium is
clean and sterile. The roots do not receive any more nutrients from a clean
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 233

medium. Some media can be reused by pasteurizing for 30 min or soaking in a

10% bleach solution for 20 min and then rinsing with tap water many times.
–– Chemical characteristics. Good cation-exchange capacity and a neutral pH (the
ability to hold nutrients).
–– Functionality. Lightweight, simple to handle, reusable, and long-lasting.

4.2.1 Rockwool

The most common fig-growing medium is Rockwool, a sterile, porous, non-

degradable material made mostly of granite and limestone that is melted and spun
into small threads before being molded into blocks, sheets, cubes, slabs, or flocks.
It absorbs water quickly and has good drainage, so it is commonly used as a starting
medium, a rooting media for cuttings, and a large biomass crop.
Advantages It has a high water retention capacity of 18% to 25% air, providing
enough oxygen to the root system if the medium is not fully submerged.
It co.mes in various sizes and forms to suit various hydroponic applications. From
1-inch cubes to massive slabs, there is something for everyone.
Mineral wool slabs can be steam-sterilized between crops and reused. However,
it takes three to four years for it to fall structurally.
Disadvantages It has a high pH, which necessitates adjusting nutritional solutions
to account for it. In addition, because the commercial material’s initial pH is high
(7.0–8.0), continual pH correction to a more suitable range (5.5–6.0) is required, or
the medium must be conditioned before soaking in a low-pH solution.
Because mineral wool does not biodegrade, it is an environmental hazard when
discarded. As a result, mineral wool has become less popular in recent years.
It has a confined root habitat and low water and nutrient buffering capacity. Even
though the water content appears high, the water passage to plant roots may be
hampered.

4.2.2 Coconut Coir

Trade names for coconut coir include Ultrapeat®, Cocopeat®, and Coco-tek®. It is
an all-natural medium comprised of shredded coconut husks. End products vary
significantly due to different suppliers and production techniques. The compressed
briquette is the most common, and it must be soaked in water before use. The coir
rehydrates during the soaking process and expands up to six times its original size.
Advantages Coconut coir is slightly acidic, holds much moisture, and provides
efficient root aeration. In addition, there are suggestions that root-promoting chemi-
cals in coir dust aid rooting.
234 W. Nosir

Coir could be used as a stand-alone medium or as part of a mixed medium for grow-
ing vegetables and cut flowers. Under mist and high humidity conditions, it can also
be used as a rooting medium for cuttings. It is biodegradable, organic, and non-
toxic, making it easy to dispose of and good for the environment.
Because it is tiny, it may be purchased compressed and inflated at home, saving
money on delivery.
Disadvantages High salinity could be a concern if the husks are soaked in saltwa-
ter during manufacture and not washed with fresh water.
Coconut coir is high in sodium and chlorine, harming plants, so it must be washed.
Calcium and magnesium are commonly added to help remove salt and provide
minerals.

4.2.3 The Aggregate of Expanded Clay

Heat dried heavy clay and stretched it to form spherical porous balls to make
expanded clay pellets. Expanded clay aggregate (ECA), grow rocks, or Hydroton®
are some of the frequent names. They are heavy enough to provide strong plant sup-
port while remaining lightweight. In addition, ECA spherical form and porosity
maintain an excellent oxygen/water balance, preventing the roots from drying out or
drowning.
Advantages Expanded clay pellets have a pH of about 7.0 and release essentially
no nutrients into the water stream. They feature a large pore space, which allows for
improved solution flow. They rarely clog or become blocked, so water drains
quickly, making them ideal for ebb and flow systems. The pellets can be rinsed and
sterilized after use and reused.

Disadvantages Compared to many other substrates, clay pellets do not have a high
water-holding capacity. As a result, they drain and dry quickly, potentially drying
out the roots.
They are somewhat pricey. In Dutch bucket systems, they frequently bond tightly
around roots and are difficult to remove. In addition, clay pellets can get sucked into
filters or drain lines and cause blockages since they float for the first few months
until they are saturated.

4.2.4 Perlite

Perlite is a naturally occurring volcanic material that expands when heated to

extremely high temperatures, becoming extremely light, porous, and absorbent. It
comes in various diameters, the most common being 0–2 mm and 1.5–3 mm. Perlite
can be used alone or in combination with other growing media.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 235

Advantages It has one of the highest percentages of oxygen retention of any grow-
ing media.
It has a high capillary activity and is highly porous. It can hold three to four times
its weight in water. Its sterility makes it ideal for germinating seeds. Root rot and
damping-off are unlikely to occur. It is reasonably priced and can be reused. It can
be steam pasteurized after use.
Disadvantages It is readily washed away due to its small weight. Because of this
flaw, perlite is an unsuitable medium for flood-and-flush hydroponic systems.
It does not retain water well when used alone in hydroponic systems such as drip
systems.
Perlite dust can cause respiratory issues and eye discomfort. Therefore it is
important to take precautions like wearing goggles and a mask when working with
it. Fans may blow it about the greenhouse once it is dry. Perlite is prone to algae
growth, which can cause problems with watering and fungus gnats.

4.2.5 Vermiculite

Is a micaceous mineral that swells into pebbles when heated to 2000 degrees
Fahrenheit. It has thought to be a tremendous rooting medium. As a result, seedlings
are frequently combined with other media types, such as coconut coir or peat moss.
It comes in various diameters, 0–2 mm, 2–4 mm, and 4–8 mm.
Advantages It has a high cation exchange capacity and can store nutrients for later
use. In addition, it is incredibly porous, has a strong capillary action, and can hold
much water.

Disadvantages When used alone, it can hold too much moisture, resulting in soggy
conditions that invite bacterial and fungal growth. It cannot be steam-sterilized
because it breaks down when heated. It is a bit pricey and may contain a small quan-
tity of asbestos.

4.2.6 Cubes of Oasis

Cubes of Oasis are water-absorbent phenolic foam, often known as floral foam,
used to make media. It is a seed and cutting growing media mostly used for plant
propagation. Oasis cubes are most commonly used for seeding lettuce and cole
crops (cabbage, collards, and kale), onions and alliums, herbs, and occasionally
tomato and eggplant seedlings.
Advantages It has a pH of 7.0 and a high water-holding capacity.
236 W. Nosir

It is a very adaptable plant that may be transferred into various hydroponic systems
and grow mediums. It is cheap, and there is no need to soak it first. It is available in
a variety of sizes.
Disadvantages There is no buffering capacity, no cation exchange capacity, and no
starting nutritional charge. It has limited utility beyond seed germination and propa-
gation. In addition, the foam may break off and clog the pump filters.

4.2.7 Sand

Sand is unquestionably the oldest and most ubiquitous hydroponic media. Other
substrates, such as vermiculite, perlite, and coconut coir, are frequently blended.
Growers frequently prefer coarse sand as a growing medium because it helps
enhance the roots’ aeration by increasing the air spaces between the grains of sand.
Advantages It is relatively affordable and widely available in most areas.
Because the finer sand particles allow lateral water circulation through capillary
action, the solution supplied to each plant is evenly distributed throughout the
root zone.
It helps aerate the mix for roots when combined with vermiculite, perlite, and
coconut coir.
Sand is highly durable because it is not chemically or biologically impacted.
It may be steam-sterilized and reused with ease.
Disadvantages It has low water and nutrient retention capacity, quickly exacerbat-
ing deficiencies. During the growth season, salt can build up in the sand. This can
be remedied by flushing the medium with purified water regularly. It is quite hefty.

4.2.8 Peat Moss

Sedges, grasses, and mosses are among the partially degraded marsh vegetation that
makes up peat. In horticultural taxonomy, sphagnum peat moss, hypnum peat moss,
and reed and sedge peat moss are the three forms of peat. The most desirable and
popular form is sphagnum peat moss, which has a higher moisture-holding capacity
and does not break down as quickly as other varieties of peat.
Advantages Peat moss has a high moisture capacity, holding up to 10 times its dry
weight in water. With a pH of 3.8–4.5, most peat mosses are acidic, which can ben-
efit some acid-loving plants. Peat moss can drain freely despite its incredible ability
to absorb water. Excess water quickly evaporates as it passes through the material.
The disposal of used peat moss does not harm the environment.
Disadvantages It is widely considered a suitable environment for various soil-
borne diseases. Although peat can be sterilized, this does not solve the problem
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 237

since sterilization creates a biological vacuum that pathogenic fungi can quickly fill.
Because its acidic nature can be detrimental to some crops, lime or dolomite is com-
monly added to raise the pH. Peat moss extraction from bogs is a disruptive opera-
tion that eliminates layers that have taken centuries to form.

4.2.9 Growstones

Recycled glass is used to make Growstones. They are thin, irregularly shaped,
porous, and reusable. They can wick water up to 4 inches above the waterline and
have good wicking ability. To keep the stems from decaying, proper drainage is
essential.
Advantages Because grow stone is inert, it provides no additional inputs or ele-
ments to the plants that could interfere with the nutrient solution in the system.
It has a high porosity and allows for many aerations of the roots.
It is non-toxic and guaranteed to be free of pollutants such as germs because it is
constructed of glass. In addition, grow stones can be reused or repurposed again.
Disadvantages They tend to hold the plant roots too tightly, so growing stones can
sometimes cause root injury. This also makes moving the plants from one medium
or grow space to another challenging.
Growstones are coated in fine silica dust that must be carefully wiped away. Because
the dust can clog drains and is dangerous to breathe, it is best to do this outside or
in a well-ventilated area.

4.2.10 Husks of Rice

Rice hulls are a rice industry waste. Even though it is organic plant material, it
degrades slowly, similar to coconut coir, making it an ideal hydroponic growing
substrate. It is frequently used as part of a growth media mix of 30–40% rice hulls
and pine bark. The different types are fresh, aged, composted, parboiled, or carbon-
ized rice hulls. As a medium modification, parboiled hulls have outperformed
other hulls.
Advantages Parboiled and composted rice hulls have an overall pH of 5.7–6.5,
which is right in the sweet spot for most hydroponically grown plants.
They have a water-holding capacity per weight comparable to perlite but a higher
air-porosity ratio and can hold more oxygen in the root zone.
They drain well and, in general, hold very little water.
Disadvantages Manganese levels in fresh and composted rice hulls are frequently
high. Manganese poisoning can occur if the pH is not correctly regulated.
238 W. Nosir

Rice hulls perform well when blended with peat or coir, but they do not work as well
as a stand-alone medium. This is because it has a poor capability for cation exchange.

4.2.11 Pine needles

Pine bark that had been composted and aged was one of the first growing media
used in hydroponics. It was once thought to be a waste product, but it has since been
helpful as a ground mulch and a substrate for hydroponically grown vegetables.
Advantages Pine bark resists decomposition better than other tree bark and con-
tains fewer organic acids that can seep into the nutrient solution.
Used bark is a naturally biodegradable substance that can be reused in various ways,
including as mulch. It keeps nutritional solution and oxygen well due to its fibrous
structure with pockets of different sizes.
Disadvantages It readily absorbs water, which can lead to flooded circumstances.
Pine bark floats, which can cause issues with ebb and flow systems. It is more suited
to a drip or wick setup. Pine bark has an acidic pH, which could be a drawback.

4.2.12 Pumice

Pumice is a volcanically formed siliceous substance. It has been graded and kiln-
dried to an internal temperature of 80 degrees Fahrenheit, making it sterile and
ready to use. Pumice media Promote aeration and drainage; it can be blended with
other forms of growing media like vermiculite or coir.
Advantages It degrades slowly and is relatively light.
Its light tint is an excellent medium for summer growing because it does not absorb
heat. In addition, it has a high level of oxygen retention.
Disadvantages It has almost identical qualities to perlite, although it does not
absorb water.
It may be too light for some hydroponics systems if purchased in little bits.

4.2.13 Sawdust

Many factors influence how sawdust performs; the most important is the type of
wood utilized and its cleanliness. The best results have been observed using sawdust
from Douglas fir and western hemlock, but western red cedar is harmful and should
never be utilized. In addition, water distributes better laterally through moderately
fine sawdust or sawdust with a higher proportion of planer shavings than coarse
sawdust.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 239

Advantages The best part about sawdust is that it is frequently inexpensive or free.
Because it holds much moisture, it must be watered with caution.
Disadvantages Sawdust may accumulate salt levels that are harmful to plants. As a
result, the sodium chloride content should be determined before using the samples.
Sawdust should be completely leached with fresh water if a substantial amount of
sodium chloride is discovered (more than ten ppm). In addition, growers must guar-
antee that their sawdust is free of soil, diseases, and chemicals from wood-processing
industries and undesired tree species.

4.2.14 Polyurethane Slabs

Polyurethane grows slabs and cubes are an uncommon hydroponics medium that
can be utilized as a beginning cubes alternative to oasis cubes or Rockwool. It is
available in the form of poly foam in hobby and fabric stores. It comes in a variety
of thicknesses and widths in rolls or sheets. Self-made starter cubes can be created
from 1- to 2-inch-thick polyfoam sheets/rolls.
Advantages It is a less expensive alternative to Rockwool or oasis cubes for seed
beginning.

Disadvantages It could include potentially dangerous substances. On the other

hand, it is unlikely to contain predefined seed germination holes.

4.2.15 Gravel

Gravel has proven very effective, especially in ebb and flow systems. It is a frag-
mented media made of rocks such as sandstone, limestone, or basalt, with huge gaps
between each particle. This provides a good supply of oxygen to the roots, but the
medium does not keep water effectively, causing the roots to dry out soon.
Advantages Gravel usually is inexpensive, works well as a beginning media, and is
relatively easy to get by. In addition, it is durable and reusable as long as it is washed
and sterilized between crops.
Its structure does not degrade, and it can be reused.
Disadvantages It is tough to handle due to its hefty weight. Gravel is not suited for
plant roots with much weight. Shale that has been expanded. Expanded shale is
formed when quarried shale is heated to temperatures above 2000 degrees
Fahrenheit. The shale is rendered chemically and physiologically inert due to the
treatment. The water in the heated shale evaporates, causing the shale to expand. It
is regarded as one of the most effective aquaponics grow media. It is light and easy
to use in aquaponic grow beds. Each stone has a wide surface area for the microor-
ganisms that convert ammonia to nitrates to thrive.
240 W. Nosir

Fig trees are unlike other deciduous fruit trees in that the shoots continue to elon-
gate as long as favorable environmental conditions for growth. Moreover, the fruit
continues to set in the axils of the new leaves throughout the growing season. For
this reason, conditions good for shoot elongation should, if possible, be maintained
throughout the summer.

4.3 Fig Cultivation

Fig should be cultivated lightly because many feeder roots are near the surface and
may be damaged if deep tillage is used. In addition, light cultivation is necessary to
break up surface crusts caused by heavy rains followed by intense sunlight.
Cultivation close to the tree’s base is difficult if low branches are allowed to develop
parallel to the ground.

4.4 Mulching

Mulches of hay, corn cobs, pine needles, sawdust, and other coarse organic matter
have been used with figs, and the results have been excellent. Mulching generally
results in excellent soil moisture, temperature, and aeration. In addition, under a hay
mulch, available mineral elements are leaching from the slowly decomposing
organic material. Mulching also helps overcome or prevent the ill effects of nema-
todes. Since some roots grow on the soil’s surface under a mulch, the mulch should
be at least 4–6 inches deep and extend from the trunk to several feet beyond the
extremities of the branches. However, good results are obtained even where the
mulch is placed only under the branches.

4.5 Irrigation

In fig cultivation, pay careful attention to soil moisture management. The roots of
most fig trees are close to the soil’s surface and can quickly dry out. In addition, figs
are particularly vulnerable to soil-borne nematodes, which eat tiny roots and limit
water flow into the tree. Apply water to the trees when the drought worsens for these
reasons. Water stress is indicated by slight leaf wilting in the afternoon.
Mulching with straw or grass clippings helps keep soil moisture consistent and
decreases weed competition for water. Due to water stress, premature fruit drop is
common in fig cultivars with no genuine seeds. When the fig tree is cultivated in
shallow soil, and the roots are nematode-infested, this problem is widespread in hot,
dry climates. In locations with poor drainage, do not overwater.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 241

This depletes the soil’s oxygen supply, causing the tree to be harmed or die.
Regular irrigation and mulching, as well as good water management, help keep
trees healthy and vigorous and decrease fruit drops. The factors influencing a fig
tree’s vulnerability to cold harm have to do with the tree’s dormancy. A mature tree
that has lost all of its leaves and is completely dormant at the first frost can survive
significantly lower temperatures than a quickly growing tree.
Inhibit irrigation in the fall to support the onset of dormancy and reduce growth.
A fully dormant fig tree can tolerate temperatures as low as 10 degrees F. Plant fig
trees along the south side of a structure in north Texas to help prevent ice damage.
Place straw mulches around the tree’s base to keep the warm soil temperature from
harming the tree’s crown during freezes. Allow time for trees or limbs to grow
before removing the frozen limbs. After that, fresh wood can be made.
Figs need a large, continuous water supply for the highest yields throughout the
summer. This is more important for Magnolia and Brown Turkey varieties, which
tend to bear continuously during the growing season, than Celeste, which produces
little fruit in late summer and early fall. Rainfall is usually sufficient in the South to
produce a good crop, even when lack of water in late summer and fall results in
discontinuing shoot elongation and fruiting. The commercial Magnolia orchards are
located where the average rainfall is high and little irrigation is used, although sup-
plemental irrigation would frequently be beneficial. The amount and frequency of
irrigation vary depending on the soil. Irrigation is required more frequently in shal-
low or moisture-retentive soils. If shoot elongation continues and leaf size is nor-
mal, moisture is sufficient. The grass beneath the grass in lawns, bushes, or trees
may wilt in the heat, while the rest of the lawn does not. This indicates the need for
water. Some fig trees grown in lawn grasses need one or more watering. From the
standpoint of survival, figs are surprisingly drought resistant. During severe
droughts, trees shed their leaves before twig injury occurs and into a drought-
induced dormancy. Subsequent rains often result in a late fall growth, which is
likely to be injured by cold weather.

4.6 Pruning

The main reasons for pruning are to produce mechanically strong bushes, open the
frontiers to sunlight and air movement, facilitate spraying and harvesting, lengthen
the harvest season, and make cultivation easier. To avoid decay, care should be taken
in pruning to cut back to a bud or lateral growing branch. Pruning differs somewhat
with the variety; however, all suckers and low-growing lateral branches should be
removed. The center should be thinned out to permit the entrance of sunlight and air.
This helps to control leaf and fruit diseases and makes spraying easier. From the
standpoint of pruning, figs grew in the South fall loosely FIG GROWING IN THE
SOUTH 19 into two categories: (1) Those that will not tolerate heavy annual prun-
ing and (2) those that can be pruned heavily and still produce large crops. The first
group includes Celeste, Green Ischia, Mission, and Hunt. Except for Hunt, these
242 W. Nosir

varieties branch freely and produce an abundance of laterals. Since breba crops are
insignificant and vigorous, the previous season’s wood is unnecessary. However, if
the branches are cut back heavily, the resulting vigorous sprouts are only lightly
productive, severely reducing yield.
For this group, only a general maintenance type pruning is needed. This includes
heading the tree back occasionally to keep it in bounds, an annual thinning out of weak
growth in the center, and removing deadwood. The Hunt variety produces long,
unbranched growth that should be headed back to make it a branch. The Celeste variety
produces a dense bush, and the inside growth should be thinned out. The second group
of varieties includes Magnolia, Brown Turkey, and Kadota. These varieties will pro-
duce good crops even if killed to the ground. Thus, they can be pruned severely and still
produce well. The main crop is produced on new wood, but an appreciable breba crop
will also be produced on the old wood. Thus, some vigorous previous season’s wood is
desirable. However, except for Kadota, dooryard trees of these varieties are best han-
dled similarly to those of the first group. Kadota does not branch freely and often fruits
excessively in relation to the available leaf surface. They were heading back all vigor-
ous growth from one-third to one-half of its length, resulting in branching and a more
favorable leaf-fruit ratio. Commercially, Magnolia trees are pruned more heavily to
facilitate harvesting and spraying than are dooryard trees. Also, the harvest season is
lengthened because the vigorous shoots from heavy pruning continue to grow longer.
A longer harvest period is essential because nearly all of the crop is canned or made
into preserves, and it cannot be processed economically in a short time.
On the other hand, heavy pruning reduces total yield considerably, and a smaller
portion of the crop is produced early in the season when prices are highest in years
past. Fig trees were pruned back annually to short stubs. Now, most growers head back
to the previous season’s growth of about one-third to one-half. This lengthens the
season appreciably but does not reduce total yield as drastically as more severe prun-
ing. The bushes are kept low by frequently cutting back to upright growing laterals
from older wood closer to the ground. Vigorous suckers arising from below the ground
level are used to replace old leaders that have become damaged (Karami et al., 2018).

4.7 Fruit Development and Growth

Crane, 1986 studied the formation and development of fruit. The fig tree’s terminal
bud opens and grows in the spring. Both vegetative and reproductive buds are pres-
ent at the same lateral position. In cultivars like ‘Mission’ and ‘Brown Turkey,’ only
one inflorescence develops into a syconium, whereas in ‘Kadota’ and ‘Calimyrna,’
both inflorescences grow into syconium. Both inflorescences at a node may emerge
in some cultivars. Comparable to other fruits, the fig syconium has three distinct
growth periods. Crane et al. (1964) reported that the initial growth stage, Stage I, is
marked by a quick increase in diameter and a slower rate of fresh and dry growth
with almost no change in sugar accumulation; you can lose weight.
Fruit diameter, dry and fresh weights, and sugar content have changed.
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 243

Over 70% of the entire dry weight and 90% of the total sugar are produced dur-
ing the growing process. During this time, the chlorophyll level in many dark culti-
vars increased. As a result, the thickness of the fruit skin reduces rapidly, and the
color of the fruit skin changes from green to bluish.
In addition, the size of the fruit grows more extensive, and the tissue softens
occurs at the end of the fig fruit‘s growth (Chessa, 1997). In common type fig fruits,
caprified and noncaprified fruit can emerge on the same branch. In most cultivars,
the first ripening period occurs within a single fruit. The second period lasts
5–6 weeks, while the third lasts 3–5 weeks. However, there are significant variances
in the life of fig cultivars. The second stage of development is called quiescence.
When it comes to cultivars like ‘Mission’. In the autumn-producing cultivars‘Siera’
or autumn Honey,’ the second phase lasts 3–4 weeks, while in the spring-producing
cultivars‘Siera’ or ‘Autumn Honey,’ the second period lasts 3–4 weeks. The fruit
harvest is shorter and lasts only 2–3 weeks since all breba fig syconia on the same
branch are at comparable developmental stages. With or without pollination, differ-
ent fig varieties can produce fruit. There is a consistent variation in nitrate levels
between persistent and nonpersistent fig varieties (El-Shazly et al., 2014). During
Stages I and II of the summer main crop figs, the average nitrate content of persis-
tent figs is three times that of nonpersistent figs. Nitrate is not discovered in nonper-
sistent fruit by Stage III. El-Shazly et al. (2014) discovered that as nitrate levels rise,
indoleacetic acid (IAA) is inhibited, implying that the difference in nitrate levels
between persistent and nonpersistent figs is due to the regulation of indoleacetic
acid oxidase. As a result, larger auxin levels are expected in persistent cultivars. In
nonpersistent Smyrna-type figs, auxin treatment enhanced fruit sets (Aljane &
Ferchini, 2009). Auxins, gibberellins, and cytokinins, among other growth regula-
tors, can cause persistence in the Smyrna-type ‘Calimyrna’ cultivar (Crane, 1986).
Auxin-induced persistent fruits take a little longer to mature than caprifled ones,
but their shape is very similar. Because they are facultatively persistent, common fig
cultivars can yield both persistent and pollinated main-crop figs. The pollinated fig
syconium produces genuine fruits, whereas the nonpollinated fig syconium has an
expanded inflorescence with several long-styled pistillate blooms. Two varieties,
‘Autumn Honey’ and ‘Brown Turkey,’ yield caprified and nonpollinated figs on the
current year’s wood. The caprified ‘Autumn Honey fruits have a darker purple skin
color and a red pulp, unlike the noncaprified fruit‘s white, pink pulp. Caprifed fruits
are often larger and have a longer shelf life (Rodov et al., 2005; Yablowicz et al.,
2005). The quantity of crops produced by fig trees directly impacts the carbohydrate
balance of the tree.
Early spring, midsummer, and late fall are the peak times for starch concentra-
tion in Smyrna-type figs, which produce a single main crop (Aljane & Ferchichi,
2006). When the shoot and breba fruit begin to form in March, there is a drop in
sugar and a parallel increase in starch concentration. Shortages of accessible carbo-
hydrates and competition between new foliage and the breba syconium can result in
syconium drop and the breba crop‘s extinction.
Applying gibberellins to the San Pedro-type ‘Nazareth’ cultivar, or nipping of
the terminal bud, temporarily stops new foliage development, allowing breba growth
and reducing competition and syconium loss (Bostan et al., 1997).
244 W. Nosir

4.8 Harvesting

The fig is a climacteric fruit that is particularly perishable and prone to physiologi-
cal disintegration. Even when stored at low temperatures, the fruit‘s postharvest life
is estimated to be between 7 and 10 days (Aljane & Ferchichi, 2006). During the
development of tissues, extensive cell wall alteration occurs (Aljane & Ferchichi,
2006). Therefore, fundamental research into the biological and physiological pro-
cesses that occur during ripening is necessary for investigating systems in which
maturation-related biological and physiological processes are implicated in posthar-
vest degradation.
To accelerate growth and ripening, ethanol is applied to fig fruits during their late
Stage 11 development. However, the receptacle tissue and the pulpy tissue of the
drupelets within mature and ripe fig fruit are visibly separate. As a result, analyzing
both tissues as a single combination may hide some cell wall modifications critical
to understanding cell wall alteration processes, Particularly during ripening.
During sequential ripening in fig fruit, Islam et al. (1997) studied the alterations
in cell wall polysaccharides that occur inside the distinct and independent tissues of
the receptacle and the pulpy drupelets. The pectic extracts included a lot of uronic
acids and a lot of neutral sugars. The uronic acid and total sugars in the drupelets
were higher than in the receptacle at the start of fig ripening. Fig fruit XETs and
EGases are part of a gene family with divergent members regulated differently dur-
ing fig fruit ripening.
The expression of these genes is influenced by ethylene and other developmental
variables, implying that numerous actions are necessary for the cooperative remod-
eling of the hemicellulose network during fig fruit softening. The scientists (Owino
et al., 2004) concluded that, like most other fruit species investigated, the gene
products of the recovered 11 cDNAs, putatively encoding cell wall-related enzymes,
are coordinated in time and amount during fig fruit production. Allow figs to ripen
completely on the tree for the best results. However, they must be plucked as soon
as they ripen to avoid spoiling by the dried fruit beetle. Microorganisms in fully ripe
fruit induce on-the-tree deterioration or souring.
Insects, particularly the dried fruit beetle, carry these organisms into the open
eye of the Fig. Daily harvests, and the removal of overripe, damaged figs can help
decrease spoiling significantly. This is especially true for kids with wide eyes.
When collecting figs, wear gloves and long sleeves to avoid skin irritation from the
fig latex.

4.9 Disease Prevention and Control

Fig rust is a severe fungus that damages the leaves of fig trees. Physopella fici is
to blame.
Small yellowish-orange dots on the leaves are the first signs of fig rust. As the
season passes, these expand somewhat and may become quite numerous. Every
9 Cultivars and Agriculture Practice of Fig (Ficus carica) 245

year, rust causes the full defoliation of many trees in the state, leaving them with a
ragged appearance. Furthermore, trees that have been defoliated early in the season
may produce new foliage sensitive to cold harm. However, defoliation does not usu-
ally occur early enough to cause fruit loss except in late-ripening types.
Neutral copper sprays are used to keep rust at bay.
A few treatments in May or early June usually keep the trees in good shape until
the fruit ripens. One or two more applications may be required during particularly
wet seasons. When the tree’s initial leaves have achieved full size, it is a good indi-
cator to start spraying. In 3–4 weeks, the second spray should be used. It is critical
to ensure that the spray substance covers all the leaves. Growing suggested types
with a closed eye, a drooping fruit feature, and fruit-splitting resistance is the first
step in preventing losses due to souring. Most fruit sourness concerns are prevented
for most of the season by pest control and the use of resistant varietals. This organ-
ism is a fungus that is mainly found in alkaline soils. The plant’s roots are killed by
this organism, causing the plant to wither and perish quickly. Unfortunately, there is
no resistant rootstock or cultivar. The only control is reconditioning the soil before
planting, which is impractical at best. This entails using a soil acidifier to com-
pletely change the soil’s pH in the area.

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Plant Science, 168(6), 1433–1441.
Chapter 10
Physiological Behaviour of Fig Tree (Ficus
carica L.) Under Different Climatic
Conditions

Aroua Ammar, Imed Ben Aissa, Faten Zaouay, Mohamed Gouiaa,

and Messaoud Mars

Abbreviations

E transpiration rate
FV/FM maximum quantum yield of PSII
gs leaf stomatal conductance
PN photosynthesis rate
RH relative humidity

1 Environmental Constraints on Fig Growth and Fruiting

Fig (Ficus carica L.) is one of the oldest fruit trees grown in the world (Mars, 2003;
Oukabli et al., 2008), especially in the Mediterranean basin and the Middle East
regions, which cover more than 90% of fresh fig production (FAOstat, 2020). Fig
growth and production are mainly dependent on climatic conditions. As limited
water resources have constantly threatened the stability of fig production, especially
in arid and semi-arid regions (Can et al., 2000), interest is increasing in evaluating
fig adaptation capacity to climatic variability. It aims to manage better water

A. Ammar (*) · F. Zaouay · M. Mars

Research Laboratory on Agrobiodiversity & Ecotoxicology (LR21AGR02), Higher
Agronomic Institute, Chott-Mariem, IRESA – University of Sousse, Sousse, Tunisia
I. Ben Aissa
Research Laboratory on Production & Protection for Sustainable Horticulture (LR21AGR03),
Regional Research Centre on Horticulture and Organic Agriculture, BP 57, 4042, Chott-
Mariem, IRESA – University of Sousse, Sousse, Tunisia
M. Gouiaa
Research Unit on Conservation and Valorization of Plant Resources (UR13AGR07), Higher
Agronomic Institute, Chott-Mariem, IRESA- University of Sousse, Sousse, Tunisia

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 247
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_10
248 A. Ammar et al.

resources, higher quality maintenance, and improved productivity. Traditionally, fig

has been mainly cultivated under rain-fed conditions (Can et al., 2008); however,
irrigation practices have improved fruit yield and quality (Melgarejo et al., 2007;
Al-Desouki et al., 2009). It has been reported that the productive orientation of the
fig orchard, timing of production, and fresh fig yield and quality are conditioned by
climatic conditions, and fig grows well in regions with intense solar radiance, hot,
dry summer, mild winters, and low relative humidity (Botti et al., 2003). However,
high temperatures could accelerate vegetative growth and fig fruit maturity and
decrease the active adult lifespan of the fig pollinator wasps (Jevanandam et al.,
2013). Under hot weather in some regions of South America and when temperatures
during winter reach more than 10 °C, dormancy and defoliation could be elimi-
nated, and figs could continuously grow and produce fruits (Flaishman et al., 2008).
The rainfall occurring in spring is useful for female fig tree growth but could nega-
tively affect fig yield due to the interference with the life cycle of the fig wasp
(Blastophaga psenes).
Furthermore, variable annual rainfall in rain-fed fig production areas leads to
yield fluctuation (Bagheri & Sepaskhah, 2014). Fruit split may result from rain, and
strong winds may cause fruit scarring (Flaishman et al., 2008). Moreover, young fig
plants are susceptible to winter frost, and the increase in relative humidity associ-
ated with low temperatures could lead to fruit quality deterioration. Indeed, fig pro-
duction is unsuitable for regions where the minimum temperatures could reach less
than −8 °C (Roger, 2002). On the other hand, the fig tree is generally grown under
marginal conditions such as drought and saline soils in some countries (Can et al.,
2008); that is why organic fertilization is recommended to improve tree vigor and
fruit quality (Mordoğan et al., 2013).

2 Drought Stress Mitigation by Different Approaches

in Rain-Fed Fig Cultivation

The irrigation scheduling in fig orchards has gained significant importance, particu-
larly in arid and semi-arid regions characterized by rainfall fluctuations and heat-
waves. Despite the importance of irrigation for fig yield and quality, few studies
were done worldwide on the quantification of fig water requirement. Previous
research confirmed the positive effect of supplemental irrigation in improving fig
tree morphological and physiological characteristics under drought conditions.
According to Roger (2002), theoretical water requirements have been estimated at
600–700 mm per year. De Sousa Andrade et al. (2014) studied the effect of drip
irrigation management on the growth and yield of the ‘Roxo de Valinhos’ fig culti-
var grown in Brazil. It has been demonstrated that fig water requirement was affected
by irrigation frequency, and mean crop coefficients Kc were about 0.71 and 0.51 for
2- and 4-days irrigation intervals, respectively.
10 Physiological Behaviour of Fig Tree (Ficus carica L.) Under Different… 249

In comparison with non-irrigated plants, irrigation increased about 27% in fruit

total yield of irrigated fig trees at a 2-days interval. Abdolahipour et al. (2019)
showed that the application of supplemental irrigation with 2000 l of water per tree
in early spring near tree trunk during drought periods is recommended to provide
high soil water status, promote vegetative growth, and increase the number and the
size of fruits for fig trees grown in rain-fed orchards. However, in the case of limited
water resources, the application of 1000 l of water per tree could also be efficient.
Al-Desouki et al. (2009) also reported that applying supplemental irrigation by
80–90 mm/ tree in the first of May, June, July, or August and using anti-transpirant
agents, namely folicote and vapor-guard, increased vegetative growth, fruit quality,
and yield of the Egyptian ‘Sultani’ fig cultivar grown under rain-fed conditions.
Honar et al. (2021) had recommended applying supplemental irrigation in March
for rain-fed fig orchards in severe drought conditions to increase soil water content,
new shoot length and diameter, number of leaves per new shoot, leaf width, and
number of fruits on new shoots, fruit diameter, and fruit yield. Also, irrigation in
August gave the best fruit yield because irrigation timing was close to the fruit rip-
ening. Besides, it has been approved that irrigation in summertime improved shoot
growth to prepare for the upcoming season’s first crop proactively. Another study on
‘Sultani’ fig cultivar grown in Egypt indicated that foliar application of ascorbic
acid and citric acid up to 300 ppm had improved vegetative growth as well as fruit
number and characteristics such as total soluble solids, fruit dimension, and fresh
and dry weights of the leaves, suggesting that foliar spray of ascorbic and citric
acids could be used to enhance water stress tolerance of fig trees (Mohamed &
El-Berry, 2021). Although the total yield and some quality characteristics were
improved through irrigation (Melgarejo et al., 2007), the excess irrigation water
must be avoided due to fig sensitivity to root rot (Tapia et al., 2003). Several addi-
tional methods to minimize drought risks in rain-fed fig orchards, including severe
pruning and mulching to reduce leaf chlorosis, mechanical weed control and trunk
whitening to enhance fruit weight (Zare et al., 2021). These horticultural practices
were effective in conserving soil moisture and reducing evapotranspiration.

3 Physiological Characteristics of Fig Grown

in Different Regions

Various fig cultivars have been described worldwide, showing an impressive physi-
ological and behavioral plasticity under different climatic conditions. In semi-arid
and arid ecosystems, the fig tree has characteristic adaptations spanning from physi-
ological (photosynthesis, stomatal conductance, transpiration rate) to morphologi-
cal (vegetative growth, leaf development) and phenological (dormancy, period of
physiological activity) features (Table 10.1). It has been reported that the seasonal
variation of ecophysiological traits may depend on various factors such as cultivar
type, age, growing area, edapho-climatic conditions, and period of measurements
(Table 10.1).
250 A. Ammar et al.

Table 10.1 Ecophysiological characteristics of different fig cultivars from different origins under
different climatic conditions
Brief description of ecophysiological
Fig type and/or cultivar Context of the study characteristics
Clones of ‘Sarilop’ Evaluation of gas exchange Leaf characteristics determining
(Smyrna type) capacity of fig grown under photosynthesis capacity (stomatal
rain-fed conditions in density, leaf area, relative water
Turkey (Kutlu et al., 2000) content, and index of leaf
succulence) were significantly
different among all clones.
The mean photosynthesis values
were higher in August than in
September, reaching a maximum of
32.15 μmol m−2 s−1. The maximum
value of E recorded in August was
22.62 mmol m−2 s−1.
‘Bursa Black’, ‘Gokalp’, The gas exchange capacity Significant differences between fig
‘Morozer’, ‘Patlican’, of 15-years-old fig cultivars cultivars were revealed.
‘Sultan Selim’, Morguz, from different origins and The transpiration rate reached its
‘Halebi’, ‘Sultani’, grown under the same maximum values at 11 am in
‘Sarilop’ rain-fed conditions in ‘Sarilop’ and ‘Sultani’.
Turkey (Can et al., 2000) The highest transpiration rates were
found in cultivars adapted to a milder
and humid climate (‘Patlican’, ‘Black
Bursa’ and ‘Sultan Selim’).
E and PN rates were significantly
higher in August compared to
September (Maximum E was 20.5
mmol m−2 s−1 for ‘Bursa Black’ and
maximum PN was 29.4 20 μmol m−2
s−1 for ‘Gokalp’).
Caprifig cultivars Measurement of mean leaf CO2 assimilation rate at 1500 mmol
photosynthesis of 30-year- photons m−2 s−1 varied among
old trees located in France caprifig cultivars from 9.4 to 13.3
(Pigé et al., 2001) μmol m−2 s−1.
Clones of ‘Sarilop’ Evaluation of seasonal and Significant differences in PN
(Smyrna type) diurnal photosynthetic and throughout the season, with
transpiration rates of fig maximum values for southern and
under semi-arid and rain-fed northern parts being in August (36.3
conditions in Turkey (Can and 22.8 μmol m−2 s−1) and
and Aksoy, 2007) September (37 and 24.8 μmol m−2
s−1) under relatively lower air
temperatures (30–33°C).
Leaf temperature was the primary
factor limiting the gas exchange
capacity of fig trees.
gs was the major control mechanism
of PN and E, particularly in the
northern parts of the trees.
Maximum gs reached 600 mmol m−2
s−1 around mid-July.
(continued)
10 Physiological Behaviour of Fig Tree (Ficus carica L.) Under Different… 251

Table 10.1 (continued)

Brief description of ecophysiological
Fig type and/or cultivar Context of the study characteristics
‘Yediveren’, ‘Banana Fig’, Determination of gas The measurements showed higher
‘Nazareth’(San Pedro) exchange capacity of photosynthetic activity in June and
‘Siyah Çiçek’ and ‘Beyaz different fig cultivars in higher transpiration rates in August
Çiçek’,‘Noire de Turkey during June (time of PN varied between 18 to 25 μmol m−2
Coromb’(Common type) harvest) and August (period s−1, E varied from 10 to 20 mmol m−2
of heaviest drought stress) s−1 and gs ranged from 300 to 400
(Can et al., 2008) mmol m−2 s−1 in June.
In August, PN varied between 15 to
20 μmol m−2 s−1, E ranged from 11 to
25 mmol m−2 s−1 and gs reached 270
to 370 mmol m−2 s−1.
Significant differences were detected
in gas exchange parameters among
fig cultivars. ‘Nazareth’, ‘Siyah
Çiçek’ and ‘Beyaz Çiçek’ were
group together; whereas ‘Banana
Fig’, ‘Noire de Coromb’ and
‘Yediveren’ comprised another
second group.
Stomatal conductance depended on
tree water status, leaf and air
temperatures, and photosynthetically
active radiation.
‘De a Libra’ (Common The gas exchange capacity PN varied during leaf expansion. PN
type) of 9-years-old fig in Spain ranged from 4μmol m−2 s−1 (13 days
(González-Rodríguez and after bud break), when leaf area was
Peters, 2009) 23% of maximum, to 20 μmol m−2
s−1 on day 51.
Maximum gs and E, at full leaf
expansion were 0.4 ± 0.01 mol m−2
s−1 and 7.6 ± 0.09 mmol m−2 s−1,
respectively.
A marked decrease in PN and gs was
registered on days 30 and 85,
correlated with low relative humidity
and high temperature.
‘Kymis’, ‘Fracasana’, Evaluation of the PN varied among cultivars and with
‘San Pedro’ and ‘Politiko’ photosynthetic activity in the season.
(white cultivars) and 5-year-old fig cultivars in ‘Vasilika Black’ had the highest
‘Vasilika Black’, ‘Vasilika Greece (Pisimisi et al., value of PN (17 μmol m−2 s−1).
Melia’ and ‘Mission’ 2012) Most of the cultivars had the highest
(black cultivars) PN values in June, coinciding with
early fruit development.
(continued)
252 A. Ammar et al.

Table 10.1 (continued)

Brief description of ecophysiological
Fig type and/or cultivar Context of the study characteristics
European fig cultivars: PN and gs reached maximum values
Investigation of the seasonal
‘Mission’ and ‘Kalamon’ changes in photosynthetic in May when leaves have reached
(Smyrna type) activity and carbohydrate maximum development and
‘Fracasana’ (San Pedro content in leaves and fruit of
increased again during fruit
type) three fig cultivars grown maturation.
under rain-fed conditions in‘Kalamon’ had significantly higher
Greece (Vemmos et al., PN (18.2 μmol m−2 s−1) than the other
2013) two cultivars, attributed to the high gs
of this cultivar (350 mmol m−2 s−1) as
to the better leaf development.
Breba crop in ‘Fracasana’ early in
the season (April) did not affect leaf
PN, so leaves are not assimilated
exporters, and the main carbohydrate
source for this early fruit growth was
that stored in shoots and other parts
of the fig tree.
‘Bither Abiadh’ (San Evaluation of seasonal The vegetative cycle of a fig tree
Pedro type) and ‘Zidi’ variation of under normal conditions ‘without
(Smyrna type) ecophysiological water stress’ comprises 4 phases of
characteristics of young growth: a phase of intense growth
potted- fig plants fully (May-June: the beginning of the
irrigated in Tunisia (Ammar summer season), a slower growth
et al., 2020a) phase (July-August), a medium
growth phase (September-October)
and “rest” phase during winter.
The ecophysiological behaviour of
fig cultivars was influenced by
climate change.
Changes in PN, gs, and chlorophyll
content were not significantly
affected by cultivar, and their
variation was attributed to year.
Maximum gs reached 435.4 and
370.6 mmol m−2 s−1 for ‘Zidi’ and
‘Bither Abiadh’ in the spring season
(81–88 days after bud break).
E rates fluctuated during all the
growing seasons and did not exceed
2.5 mmol m−2 s−1 for both cultivars.
The lowest values of PN and gs were
recorded in summer (156 days after
bud break) and at the end of the
growing season in autumn.
E (mmol m−2 s−1): Transpiration rate, PN (μmol m−2 s−1): Photosynthesis rate, gs (mmol m−2 s−1):
Leaf stomatal conductance
10 Physiological Behaviour of Fig Tree (Ficus carica L.) Under Different… 253

4 Fig Responses and Resilience Towards Drought Under

Different Climatic Conditions

To help predict future impacts of climate change on fig tree growth and yield, more
and more profound knowledge of the ecophysiological mechanisms of the fig tree is
needed. Simple stress indicators based on easily measurable traits are needed to
overcome this. Different physiological behaviors under water stress have been
observed in fig; according to the cultivar, the growing conditions, and the degree of
stress imposed on fig. Genotype related differences have been described in fig
responses to drought in terms of vegetative growth (El-Shazly et al., 2014; Rostami
& Rahemi, 2013a; Shahidi-Rad et al., 2015), fruit production (Jafari et al., 2012),
biochemical changes including increased activity of radical scavenging enzymes
such as peroxidase (POD), superoxide dismutase (SOD), ascorbic peroxidase
(APX) and catalase (CAT), osmolytes accumulation in particular proline (Shirbani
et al. 2013; Rostami & Rahemi, 2013b; Shahidi-Rad et al., 2015) and producing
proteolytic enzymes such as chlorophyllase (Gholami et al., 2012). Some different
cultivars may behave similarly. For example, Ammar et al. (2020b) found that ‘Zidi’
as Smyrna type and ‘Bither Abiadh’ as San Pedro type exhibited the same response
to water stress and recovery.
Studies aiming to understand fig tree gas exchange capacity in response to water
deficit are not frequent. However, Ammar et al. (2020b) demonstrated that stomatal
control of gas exchange was crucial for fig drought response. Indeed, the reduction
of stomatal conductance has been considered an immediate response to water stress
before any change was detectable in leaf water content (Pirasteh-Anosheh et al.,
2016), showing the important role played by abscisic acid (ABA) in promoting
stomatal closure (Bondada & Shutthanandan, 2012).
To cope with water stress, fig trees developed two different strategies to overcome
the dry and hot summer period and complete their life cycle. In this sense, an adap-
tive strategy relies on maintaining low rates of active photosynthesis and reducing
water loss. This took place when moderate water stress was applied by reducing fig
water requirements by 40 and 60% (Caruso et al., 2017) or by 50% (Ammar, 2020),
which induced a decrease in stomatal conductance (less than 100 mmol m−2 s−1)
associated with a reduction of photosynthesis rates. Besides, moderate water stress
also affected the plant water status by decreasing leaf relative water content and
transpiration rate and an increase of leaf temperature owing to stomatal closure, with
not very high rates of leaf senescence. Furthermore, regarding fig fruit quality, regu-
lated deficit irrigation with 55% ETc significantly reduced fruit skin side cracking
and ostiole-end splitting at harvest, improving marketable yield (Kong et al., 2013).
Prolonged drought by suspending irrigation for 2 weeks (Ammar et al., 2020b)
or applying prolonged moderate water stress by reducing to 50% the water supply
combined with high temperatures exceeding 45 °C (Ammar, 2020) induced a sharp
decrease in stomatal conductance, photosynthesis activity, transpiration rate and
maximum quantum yield of the photosystem II (PSII), chlorophyll degradation and
necrotic spots in leaves, resulting in growth inhibition and leaf abscission. This cor-
responded to an avoidance strategy to overcome drought periods associated with
extreme heat stress during summer. This pattern could be considered ‘summer rest’
254 A. Ammar et al.

when the fig tree exhibits a temporary growth suspension and massive leaf abscis-
sion to evade the unfavorable hot and dry season (Shahidi-Rad et al., 2015; Ammar
et al., 2020b).
Fig has revealed ‘drought stress memorizing’ by showing a rapid re-growth after
subsequent re-irrigation, extending the annual vegetative cycle and biomass accu-
mulation. Indeed, plants could survive after leaf fall by keeping the stem alive dur-
ing the drought period and accumulating carbohydrates in the stem and roots to
stimulate the emergence of new leaves once the stress is relieved (Xu et al., 2008,
2012; Naidoo & Naidoo, 2018). Furthermore, plant recovery after re-irrigation
induced an increase in fructose in leaves related to a decrease of sucrose in leaves
and stems, suggesting sucrose cleavage and mobilization for the growing buds after
leaf abscission (Xu et al., 2008).
Fig plants, which were more affected by high air temperatures combined with
water stress, required less time to resume vegetative development (Ammar, 2020).
Furthermore, new leaves of re-watered fig plants showed high stomatal conductance
reaching more than 300 mmol m−2 s−1, associated with high photosynthesis activity
and chlorophyll content compared to leaves of non-stressed plants. Also, water
stress recovery resulted in an extended vegetative growth until December (Ammar
et al., 2020b). The cycle length of the fig could vary according to the genotype, solar
radiation, accumulated degree-days, and air temperature as well (Ferraz et al., 2020).
Thus, the strategy to cope with water scarcity is mainly oriented toward enduring
the summer period under different climatic conditions, as summarized in Fig. 10.1.

Fig. 10.1 Fig ecophysiological responses and strategies to water stress and recovery
(–): negative effect, (+): positive effect
10 Physiological Behaviour of Fig Tree (Ficus carica L.) Under Different… 255

Regardless of water stress intensity and duration, fig could be classed as a drought-
stress resilient species considering its rapid growth recovery. In this sense, the inter-
est in understanding fig physiological control of abiotic stresses has increased.
Nevertheless, most published research was conducted on potted-fig plants and cov-
ered just a short period of exposure to water stress in experimental conditions,
which is not comparable to the real field conditions. Furthermore, as fig represents
a vast genetic diversity (Moniruzzaman et al., 2020), more effort is needed better to
understand different fig cultivars‘behavior under field conditions and identify the
most suitable cultivars for cultivation in a given region.

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Chapter 11
Fig (Ficus carica L.) Production and Yield

Malek Ben Temessek, Wafa Talbi, Hana Chrifa, and Sami Fattouch

Abbreviations

CEC Cation Exchange Capacity

GA Gibberellic acid
Kg ha Kilogram per hectare
SSC Soluble solid concentration
TA Titratable acidity
TSS Total soluble solid

1 Conventional and Molecular Breeding Systems in Fig

(Ficus carica L.)

The fig tree (Ficus carica L.) is a diploid heterozygous (2n = 26) plant. The DNA
2C levels are quite low, ranging from 1.37 pg to 1.37 pg, corresponding to 270 mil-
lion base pairs and 50,000 genes. People from Persia, Asia Minor, and Syria spread
the fig tree throughout the Mediterranean region. It has been an essential food cul-
ture for thousands of years, and it is considered highly good in the Mediterranean
diet (Solomon et al., 2006). Thousands of cultivars, most unnamed, have been
developed or introduced, and the fig has spread far beyond its natural area due to
human migration. Most Mediterranean countries, Europe, North, South Africa,
America, and many other Asian countries now grow them. The monoic ancestor of

W. Talbi
M. Ben Temessek (*) · H. Chrifa · S. Fattouch
EcoChem Laboratory, Department of Biological and Chemical Engineering, National
Institute of Applied Sciences and Technology (INSAT), University of Carthage (UCAR),
Tunis, Tunisia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 259
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_11
260 M. Ben Temessek et al.

the common fig tree (Machado et al., 2001) evolved into gynodio species with
bisexual (functioning) male figs or caprifigs and unisexual female trees. Female and
male flowers are arranged inside the syconium, a constructed fig fruit structure.
Hundreds of female flowers are included in a female fruit. A male fig plant, called
caprifig, has hundreds of separated male (staminate) flowers and female (pistillate)
flowers. The differentiation between the hermaphroditic and female strains forms
establishes the basis for maintaining the close symbiotic relationship between Ficus
plants and the Blastophaga wasps (Galil & Neeman, 1977).
Plant breeding is a dynamic area of applied horticultural science. It relies on
genetic variation and uses breeding to enhance traits and characteristics of interest
to growers and consumers. The genus Ficus has two reproductive systems, the
monocecity (approx. 400 species) and gynodioecy (350 species, including Ficus
carica L.) (Berg, 1989). In Ficus, monoecy is hypothesized to be ancestral, and
gynodioecy derived. In monoecious figs, each fruit (syconium) has both male and
female florets functioning to produce pollen and viable seed. Some florets are para-
sitized to produce wasps that carry the pollen to other trees. In gynodioecious figs,
separate trees produce syconia creating either viable seed or pollen and wasps. The
studies of fig floral biology and the understanding of the coevolution with the fig
allowed the establishment of fig breeding techniques (Weiblen, 2002). The juvenile
stage can be reduced by grafting seedling buds into older trees or pumping and
anchoring them in a vertical position at the beginning of development (Stover et al.,
2007) and then growing them intensively, using the last method under the warm
climate conditions of Israel. Indeed, Flaishman et al. (2008) obtained reproductive
development and performed fruit evaluation as early as one year after seed extrac-
tion. The majority of today’s commercial fig varieties are thought to be the product
of unintentional selections made by unknown growers. Local fig collections exist in
several places, used to select superior clones (Mars et al., 1997; Mars et al., 2009).
Due to its very narrow industry, fig breeding projects have a restricted budget and a
small number of breeding programs.
In the United States, two breeding programs were conducted at the beginning of
the twentieth century; the largest was at the University of California, led by
I.J. Condit and Storey (begun by R.E. Smith in 1922). At the same time, Slykov led
another breeding program in the Soviet Union. Doyle and Ferguson (1997) contin-
ued the fig breeding program in California. In addition, fig crosses and the heritage
of various fig features have been described in many countries like Japan (Awamura
et al., 1996, 1997), Turkey, and other nations.

2 Fig Production, Research, and Development

Fig fruits are transformed into several processing products, such as dried figs, pre-
served fruits, jam, juice, wine, powder, and others, but the most popular ones are
preserved fruits and jam. There are nearly 20 figs-processing factories on the
11 Fig (Ficus carica L.) Production and Yield 261

mainland. Fig fruits and leaves contain many amino acids and inorganic elements,
showing their high nutritional value. Furthermore, fig contains several medical
components such as flavones, rutin, and quercetin, which could be used in cardio-
vascular disease medicine production. Additionally, the cooperative research
showed that the extracts from fig had significant inhibition in the growth of several
cancer cells in mouse bodies, the inhibition for spleen cancer high up to 64.0%,
EAC 53.8%, Lewis lung cancer 48.8%, HAC liver cancer 44.4% and S180 cancer
41.8%. Furthermore, the nontoxic extracts of figs could increase the body’s immu-
nity and postpone the life span (Arpaci, 2015).

3 Fig Fruit Latex Yield and Protease Activity as Influenced

by Cultivar, Maturity Stage and Harvesting Time

Laticiferous cells generate fig latex, commonly coupled with other botanical com-
ponents like a blue flag, barley, fenugreek, ginger, and hot pepper to soften solid
tumors and treat skin problems (Lansky et al., 2008). Most species’ latex contains a
wide range of physiologically active chemicals. Many of these chemicals provide
herbivore resistance through toxicity or antinutritive effects, while others contribute
to the stickiness that can ensnare insect herbivores (Lazreg-Aref et al., 2011).
Several defense-related components (e.g., rubber, cysteine protease, alkaloids)
appear in the latex of distant phylogenetic groups, suggesting common functions
and convergent evolution (Agrawal & Konno, 2009). The volatile compounds
released by fig latex comprise mainly mono- and sesquiterpenes (Hossaert-McKey
et al., 2010). Oliveira et al. (2010) identified 34 compounds distributed by distinct
chemical classes: five aldehydes, seven alcohols, one ketone, nine monoterpenes,
nine sesquiterpenes, and three other compounds. Moreover, thirty-six coumarins
were identified from Tunisian caprifig latex, which caused a strong bactericidal
effect (Lazreg-Aref et al., 2012). The volatile compounds are critical factors in the
fig-fig wasp mutualism. Indeed, each wasp species seems to be attracted to specific
chemical compounds of its associated fig tree species (Chen & Song, 2008; Chen
et al., 2009). Maffei (2010) demonstrated the involvement of the volatile organic
compounds in the attraction of pollinating and predatory insects.
In some cases, pollinators of some species are stimulated by the emitted volatile
compounds of their associated fig species and generally not by the volatile com-
pounds of another species (Grison-Pigé et al., 2002). Fig latex is involved in defense
mechanisms by protecting ripening fruits against plant pathogens like fungi and
insects, which they can attack directly (Konno et al., 2004). There was a substantial
difference in latex yield for the several fig cultivars between harvesting time and the
fruit maturity process. The highest values were achieved in the mid maturity stage
for all cultivars, with no significant changes between stages (Table 11.1).
262 M. Ben Temessek et al.

Table 11.1 Protease activity and latex yield of Smyrna-type fig cultivars during three times a day
Latex Protease Latex Latex Protease Latex
Maturity harvesting activity yield Maturity harvesting activity yield
Cultivar stage time (U/ml) (ml) Cultivar stage time (U/ml) (ml)
Khedri Unripe 1 324685 a
0.21b
Bidhi Unripe 1 523363b 0.20NS
2 204671 b
0.22b
2 556677 ab
0.00NS
3 419712a 0.44a 3 598765c 0.40NS
Mid- 1 229760a 0.50b Mid- 1 191212a 0.50NS
maturity 2 303214b 0.49b maturity 2 102312b 0.50NS
3 330329a 0.80a 3 113321b 0.80NS
Ripe 1 11563c 0.10 Ripe 1 89754a 0.12b
2 102866b 0.10 2 34254c 0.10b
3 228427a 0.10 3 66654b 0.20a
Chetoui Unripe 1 1,008,181 0.10NS Zidi Unripe 1 6940901b 0.10NS
2 201,315 0.10NS 2 443312c 0.10NS
3 1,141,515 0.10NS 3 798865a 0,30 NS
Mid- 1 131,567 0.20NS
Mid- 1 988076 a
0,30 NS
maturity 2 123,899 0.10NS maturity 2 808765 b
0,30 NS
3 1,107,181 0.40NS
3 1012345 a
0,60 NS
Ripe 1 1205181b 0.10NS
Ripe 1 990865 a
0,20 NS
2 123,498 0.05NS
2 809743 b
0,17 NS
3 191,418 0.20NS
3 988065 a
0,30 NS
Soltani Unripe 1 634090 a
0.70NS
Delgane Unripe 1 580909 b
0.20NS
Abiadh 2 609854 ab
0.30NS
2 446785 c
0.10NS
3 568743 b
0.30NS
3 665432 a
0.40NS
Mid- 1 556698 b
0.50NS
Mid- 1 764367 b
0.50NS
maturity 2 412211 c
0.40NS maturity 2 708743 b
0.30NS
3 709854 a
0.90NS
3 806534a 0.70NS
Ripe 1 209812b 0.35NS
Ripe 1 664365 a
0.10NS
2 123421a 0.20NS
2 609865 a
0.10NS
3 223254a 0.40NS 3 667898a 0.20NS
Results are expressed as means ± deviation (n = 3). Different small letters within one cultivar are
significantly (P < 0.05) different according to Duncan test. Capital letters within cultivars are sig-
nificantly different (P < 0.05) according to Duncan test. *Significant at P < 0.05. NS non-significant.
(1: morning, 2: mid-day and 3: late afternoon) afternoon). (Lazreg-Aref et al., 2018)

4 Fig Production in Subtropical South-Western

Saudi Arabia

Fig has been considered an essential crop for the people of southwest Asia for quite
a long time. Persians, Arabs, Canaanites, and Egyptians cultivated figs for more
than 3500 years (Obenauf et al., 1978). Despite being cultivated in this part of the
world for millennia, fig Production remains limited in Saudi Arabia. Two main cul-
tivars of figs could be found in the southwestern part of the Kingdom, ‘Baladi’ and
‘Brown Turkey’. Both cultivars are of the common type. The first is a local cultivar
11 Fig (Ficus carica L.) Production and Yield 263

with small sweet fruit a yellowish-green skin. The latter, introduced in the early
eighties, has a medium-size fruit with brownish skin. The fruit matures in late sum-
mer at high elevation (Assir) and winter (December–February) (off-season) in the
plains and foot hills. Fruit quality in Assir is good, but farm size is limited by the
lack of water for irrigation. Therefore, most acreage is concentrated in the coastal
region, where water for irrigation is available. The Saudi Ministry of Agriculture
does not publish separate statistics for fig production or acreage.
However, the terminal report of the FAO/Saudi Arabian Government Cooperation
Program (UTFN/SAU.013/SAU) gives an estimate for the number of fig trees in the
Jazan province alone to be about 311,300 (FAO, 2007). However, heavy tree losses
to termites and diseases were reported by growers during this last decade.
Nevertheless, a sizable crop is produced with small quantities (150–200 t) of fresh
fruit exported to Europe and Japan. Presently, the crop is mainly sold locally for
reasonable prices; no drying or canning is practiced. However, as new plantings
come to bearing, such infrastructure will be needed to maintain orchard profitabil-
ity. Furthermore, the lack of an efficient marketing strategy thwarting fig production
progress despite the region’s enormous potential.

5 Fig Pollination and Caprification System

The fig tree (Ficus carica L.) is a functionally dioecious species. The male fig tree
or caprifig ensures the pollen production and survival of the symbiotic pollinator,
blastophagus (Blastophaga psenes L.). The blastophage reproduces exclusively in
caprifig. Female or domestic fig trees produce figs pollinated in July and ripe in
autumn: these are the autumn figs. A few fig buds do not develop during the summer
but only begin to grow the following spring.
On most domestic figs, these figs fall before the stage of pollen receptivity; it is
the fall early. Nevertheless, some particular genotypes retain figs to receptivity. At
this stage, the figs cannot be pollinated because there is no pollen available at this
time of year, but they usually continue to develop by parthenocarpic way to give ripe
figs in July; fig flowers. These fig trees are called bifères as opposed to fig trees that
lose their flower figs before receptivity, said unifying. Much has been written about
the extraordinary complexity of the biological reproduction of Ficus species, par-
ticularly regarding pollination (Dickson & Dickson, 1996). Ficus species are gyno-
dious and functionally dioecious. Some of them are functionally female and produce
only one fruit seed, while others are functionally male and produce only pollen and
pollen wasp offspring (pollen carrier); it is then Blastophaga psenes that brings the
pollen of the male flower to the female flower (Janzen, 1979; Weibes, 1979). On
figs, receptivity is essential in developing the fruit distinguished by volatile emis-
sions to attract pollinators.
Pollination is essential for proper fruit development (Wagner et al., 1999).
Blastophaga wasp and caprifig are necessary for caprification and normal fruit
development. If this fertilization process does not occur, the fruits do not develop
264 M. Ben Temessek et al.

properly and will fall from the tree. Many fig cultivars are totally or partially parthe-
nocarps; the synconium swells without pollination and does not have only underde-
veloped infertile seeds (Dickson & Dickson, 1996). Pollinated figs are generally
larger and greener than unpolluted figs, with a darker pulp (Oukabli et al., 2003). In
fig orchards, for high economic return, the pollination process must be repeated two
or three times as the figs’ synconium gradually becomes receptive. Thus, it is essen-
tial to have two or three-period cultivars (Zare et al., 2013). The number of spent
caprifigs and the pollination period depend on the female cultivar‘s weather condi-
tions, age, size, and yield. The common practice of caprification is to distribute the
male figs at intervals of a few days over approximately 3 weeks (Rahemi & Jafari,
2005; Zare et al., 2013).

6 Micropropagation of Ficus carica L. Studies

on the Propagation of Fig Cultivars by Budding

In India, figs are considered small commercial fruit because their output cannot
compete with other commercial fruit crops. The major causes for insufficient yield
include a lack of knowledge on varietal introduction and characterization and a
scarcity of high-quality planting material. Tip cuttings are used to grow fig trees
(Dolgun & Tekintas, 2008; Lionakis, 1995; Mansour, 1995; Kai et al., 1997). Fig
cuttings usually are simple to root; however rapid variations in air temperature and
soil moisture conditions impair rooting and shoot growth. Furthermore, soil-borne
bacterial and fungal infections impact production cost and quantity during growth
in the field (Kabasakal, 1990).
Consequently, nursery plant production efficiency in the field is around 50–60%
(Cobanoglu et al., 2004). Cuttings are frequently used to propagate fig plants
(Dolgun & Tekintas, 2008). However, fewer nursery plants are created because of
the restricted branches generated by traditional layering and bottom shoots proce-
dures. Thus, these approaches are not commonly used in commercial nursery plant
production (Dolgun & Tekintas, 2008).
Because of the constraints above in fig nursery plants, intended success and sus-
tainability cannot be guaranteed. Furthermore, 5–10 cm long shoots are abundant;
however, they are unsuitable. However, these shoots can be used for budding with
success. New fig cultivars have been introduced at the Punjab Agricultural University
in Ludhiana. Some of them show great promise in terms of yield and fruit quality.
However, the introduction of fig as a new crop in Punjab necessitates many plants,
which may be difficult to meet using traditional techniques (Rattanpal et al., 2014).
Furthermore, several potential genotypes grow poorly on their roots under Punjab
circumstances. As a result, the ability to swiftly and cheaply produce fig plants
would have significant commercial value.
Thus, the research conducted at the College Orchard of Punjab Agricultural
University was based on analyzing thirteen fig tree cultivars obtained from various
11 Fig (Ficus carica L.) Production and Yield 265

Indian and alien sources for their capacity to be propagated by the budding method.
The thirteen cultivars of fig (Ficus carica L.): ‘Black Fig’, ‘Brown Turkey’,
‘California Brown Turkey’, ‘Conadria’, ‘Deanna’, ‘Dinkar’, ‘Genoa’, ‘Golden
Celeste’, ‘King’, ‘Panache’, ‘Poona’, ‘San Piero’ and ‘Texas’, were budded on
‘Brown Turkey’ seedlings, during the last week of August in 2012 and 2013. The
cultivar ‘Brown Turkey’ had the highest mean budding success (93.4%), while
‘Black fig’ had the lowest (70.3%) (Rattanpal et al., 2015). There was no significant
difference between the cultivars regarding the number of days it took from budding
to sprouting. Furthermore, no incompatibility was found in any cultivar even 2 years
after budding. The union of most cultivars got smoother with time.
Moreover, none of the plants split from the bud union; instead, the union
smoothed out over time. This method not only saved the propagules, but prelimi-
nary investigations suggested that budded plants were more vigorous than cuttings-
propagated plants. The cultivar ‘Genoa’ plants were far more robust on ‘Brown
Turkey’ rootstock than on their roots. This is verified by the findings of Krezdorn
and Glasgow (1970), who budded or grafted various F. carica cultivars onto F. glom-
erata, F. cocculifolia, F. graphalocarpa, and F. palmate and found no incompatibil-
ity with any of the rootstock species. Similarly, in Japan, Hosomi et al. (2002) used
rootstock to gain an advantage in fig varieties by grafting sensitive cultivars on
resistant rootstocks. Although fig culture in Indian Punjab is new, this approach will
facilitate the rapid dissemination of cultivars discovered to be appropriate for Punjab
conditions (Rattanpal et al., 2015).

7 Fertigation with Potassium Increases Size and Yield

in Fresh Figs

California is the major producer of figs (Ficus carica L.) in the United States, and
the state’s fresh fig industry has lately expanded. In addition, several countries have
reported the benefits of nitrogen (N), phosphorus (P), and potassium (K) fertiliza-
tion with additional calcium (Ca) or K treatments during the growing season,
increasing overall quality by reducing the number of fruit with ostiole-end cracking
and sunscald on ‘Calimyrna’ or ‘Sarlop’ figs (İrget et al., 2008), and others (Antunes
et al., 2008; Irfan et al., 2013).
Indeed, studies carried out by Holstein et al. (2015) conducted two seasons of
research to assess the effects of K and Ca, both foliar and fertigation treatments, on
fresh fig fruit size, yield, and soundness. Using a foliar spray containing K and Ca
to ‘Black Mission,’ ‘Brown Turkey,’ and ‘Sierra’ decreased side cracking and
enhanced fruit size. The response to foliar treatments, on the other hand, was very
varied amongst plants, suggesting that absorption issues exist. Furthermore, in
‘Sierra’ figs, an increase in the percentage of fruit with ostiole-end splitting was
found. Fertigation with K and Ca enhanced yield in both cultivars. In ‘Black
Mission,’ the percentage of culls was 49% for control trees and 43% for treated
266 M. Ben Temessek et al.

trees. Similar findings were achieved in ‘Sierra,’ with 42% for control trees and 36%
for treated trees. As a result, marketable production was greater for treated trees
than for control trees for both varieties. In ‘Sierra,’ only fruit size was enhanced by
fertigation (40.4% compared with control 38.7%), but soluble solids concentration
(SSC) and titratable acidity (TA) were unaffected in either cultivar.
In terms of leaf macronutrient concentrations, nutritional development was com-
parable in both cultivars, with leaf N, P, and K levels dropping from late spring
(May) through fall (October) (Tables 11.2). These low N and K leaf levels in control
plants late in the season point to the possible advantages of spraying N and K during
the season. On the other hand, Ca levels increased a little late in the season. After
the June sampling, all leaf nutrients hit a plateau, suggesting that leaf sampling was
done in midsummer (July–September) will be more stable and representative than
earlier leaf sampling, as previously advocated for California dry fig orchards. Except
for late in the season for ‘Black Mission’, leaf sample nutrients revealed no differ-
ences between trees treated and controlled with K and Ca (Holstein et al., 2015).
This study found that dramatically increasing K and Ca fertigation may greatly
boost marketable production of the fresh fig cultivars ‘Black Mission’ and ‘Sierra,’
which is critical for fresh fig growers since marketable yield determines income. In
‘Sierra’ figs, irrigation treatment boosted fruit size, linked to high leaf K
concentrations.
As well as, Soliman et al. (2018) investigated the effects of potassium fertiliza-
tion on the fruit quality of figs grown in Riyadh. The findings of this study show that
potassium fertilization treatment had a substantial impact on the enhancement of the
physical qualities of fruits in both seasons (during the 2015 and 2016 seasons). The
maximum fruit quality was achieved by applying K2O at 400 g per tree, followed by
200 g per tree. The findings are consistent with Kassem (2012), who found that
potassium fertilization enhanced the fruit‘s weight, volume, and dimensions.

Table 11.2 Mean macronutrient concentrations (% dry weight) of ‘Black Mission’(a) fig and
‘Sierrra’ (b) fig leaves collected on four sampling dates from trees fertigated with 12.6 units of K
ha−1 and 2.5 units of Ca ha−1 grown at the Kearney Agricultural Research and Extension Center,
Parlier, CA. Each mean is a composite sample of 10 trees
Control Treated Control Treated Control Treated Control Treated
Date N N P P K K Ca Ca
(a) ‘Black mission’ fig treatment
May 12 2.89 2.99 0.156 0.164 1.27 1.36 2.40 2.24
Jun 29 2.24 2.30 0.119 0.123 1.23 1.35 3.12 2.90
Aug 10 2.14 2.21 0.122 0.115 1.21 1.20 2.46 2.94
Sep 11 1.98 2.13 0.109 0.109 1.08 1.42 3.33 3.32
(b) ‘Sierra’ fig treatment
May 12 3.21 3.15 0.166 0.172 1.64 1.44 2.79 2.49
Jun 29 2.34 2.41 0.136 0.133 1.79 1.88 3.35 3.41
Aug 10 2.20 2.00 0.132 0.114 1.65 1.54 3.21 3.51
Sep 11 2.13 2.09 0.118 0.107 1.66 1.52 3.50 3.80
Holstein et al. ( 2015)
11 Fig (Ficus carica L.) Production and Yield 267

According to Ganeshamurthy et al. (2011), high K concentrations are required to

obtain the largest overall fruit yield of grapes and bananas with good maintaining
quality and marketability. Thus, potassium (K) has a tight interaction with water and
plays an active part in cell swelling and expansion. More than one study found that
K supplementation increased the volume of fruits and the thickness of the crust in
citrus (Mattos et al., 2005).
Furthermore, potassium plays a vital function in soil pH adjustment, osmotic
regulation, protein synthesis, stomatal movement, photosynthesis, and cell expan-
sion (Läuchli & Pflüger, 1980). As a result, they demonstrated that potassium fertil-
ization was the most successful fertilization method for fig. Furthermore, it improved
fruits‘physical and chemical quality by increasing their weight, size, dimensions,
total sugars, reducing and non-reducing sugars, total soluble solids, and leaf mineral
content (Soliman et al., 2018).

8 Effect of Gibberellic Acid on Growth, Yield and Fruit

Quality of Fig (Ficus carica L.)

Because of its drought tolerance and simplicity of control, fig cultivation is becom-
ing more popular in arid areas, where it may be grown as a monoculture or in a
mixed crop with other fruit trees. Its incorporation into the agricultural land use
system aids crop diversity (Kurubar et al., 2017). Commercial products such as fig
syrups, jam, jelly, and spiced or pickled figs are made. Fresh or dried fig fruit is
prized for its laxative properties. Its latex is used to coagulate milk in various coun-
tries. However, the fig fruit is fragile and perishable by nature; therefore, there is a
risk of rotting after harvest.
Aside from nutrients, growth regulators have been shown to have favorable
impacts on various fruit crops. Scientists have reported that different plant growth
substances such as Parachlorophenoxy acetic acid (PCPA), indole-3-n-butyric acid
(IBA), naphthalene acetic acid (NAA), 2,4,5-trichlorophenoxy acetic acid (2,4,5-T),
gibberellic acid (GA), ethylene (C2H4), 2-(chloroethyl) phosphonic acid (ethephon)
increased yield and quality while decreasing maturity period (Kurubar et al., 2017).
The studies carried out by Kurubar et al. (2017) were based on the evaluation of the
effect of different concentrations of GA with an increased number of spray applica-
tions at vegetative bud initiation after annual pruning, 15 days after bud initiation,
and 30 days after bud initiation. It is inferred that increasing the concentration of
GA with increasing the number of sprays led to increased shoot length (2.95 cm in
Treatment 9) (Table 11.3). Similarly, for the number of functional leaves (616.2)
and the length of the fruit (5.72 cm), the behavior is reversed for the diameter, fruit
weight, and number of fruits per plant. Increased GA concentration and spray fre-
quency reduce the fruit diameter and weight, whereas the fruit diameter and weight
increase where GA concentration is lower.
268

Table 11.3 Effect of treatments with gibberellic acid (GA) on vegetative and fruit characters, quality, and number of days taken for maturity and fruit
yield of fig
Average of 2 years
Fruit Total Non
Shoot Number of Fruit Fruit Fruit Number yield Fruit soluble Titratable Reducing reducing Total
length functional Number length diameter weight of days to (kg yield solids acidity sugars sugars sugars
Treatment (cm) leaves of fruits (cm) (cm) (g) maturity plant-1) (t ha−1) (%) (%) (%) (%) (%)
T1 2.28 477.87 322.08 5.22 4.48 39.23 108.87 12.63 8.40 16.21 0.198 14.59 1.96 16.54
T2 2.40 521.43 329.36 5.48 4.13 38.10 105.80 12.54 8.35 16.50 0.203 14.79 1.86 16.64
T3 2.49 555.88 357.56 5.59 4.06 36.68 102.21 13.11 8.73 16.90 0.212 15.10 1.80 16.90
T4 2.39 516.52 335.77 5.27 4.18 39.23 108.37 13.17 8.76 16.38 0.202 15.00 1.84 16.84
T5 2.48 554.84 353.52 5.47 4.04 37.72 99.99 13.38 8.91 16.97 0.215 15.43 1.78 17.20
T6 2.50 577.43 352.86 5.62 3.94 36.98 97.79 13.09 8.71 17.27 0.230 15.57 1.72 17.29
T7 2.70 601.53 361.15 5.54 4.06 36.76 93.46 13.27 8.83 17.08 0.212 15.23 1.83 17.06
T8 2.80 600.22 360.71 5.69 3.90 36.71 93.63 13.24 8.81 17.18 0.231 15.69 1.72 17.40
T9 2.95 616.23 351.11 5.72 3.93 36.81 91.23 12.92 8.60 17.28 0.230 15.78 1.62 17.40
T10 2.22 462.62 314.73 5.08 4.67 39.26 125.83 12.35 8.22 17.39 0.196 16.02 1.98 18.00
Kurubar et al. (2017)
T1: GA 20 ppm at bud initiation (BI) stage; T2: GA 40 ppm at BI stage; T3: GA 60 ppm at BI stage; T4: GA 20 ppm at BI stage and 15 days after bud initiation
(DABI); T5: GA 40 ppm at BI stage and at 15 DABI; T6: GA 60 ppm at BI stage and at 15 DABI; T7: GA 20 ppm at BI +15 DABI +30 DABI; T8: GA 40 ppm
at BI +15 DABI +30 DABI; T9: GA 60 ppm at BI +15 DABI +30 DABI; T10: Control
M. Ben Temessek et al.
11 Fig (Ficus carica L.) Production and Yield 269

This is due to GA, which plays a key role in shoot elongation, breaks dormancy,
and utilizes carbohydrates for shoot growth rather than fruit development, and a
larger concentration of GA accelerates the initiation of leaf formation (Gaikwad,
1975). T9 demonstrated that sprays’ enhanced concentration and quantity resulted
in early fruit maturity. The fruit production per plant and hectare showed a non-
significant relationship between GA content and spray frequency. Among the differ-
ent concentrations of GA spray-on fig, the highest rate for yield per plant and yield
per hacter were noted in T5 (13.3 kg and 8.91 t ha−1, respectively). Thus, the total
soluble solid (TSS), acidity, reducing sugars, and total sugars, were improved with
the addition of GA concentration and frequency of sprays, but, in the case of non-
reducing sugars, the effect was reversed. However, the control treatment outper-
formed the chemical spray in statistical significance.
The rise in concentration causes an increase in fruit length, which has no eco-
nomic benefit because it negatively influences fruit weight, which is a significant
contributing yield characteristic. Therefore, GA spray may not be helpful unless
paired with other growth regulators that might cause fruit thickness after elongation.

9 Increasing the Tolerance of Fig Trees to Drought Stress by

Trunk Thinning

The fig tree is indigenous to Iran, East Asia, and Syria. One of the most serious
issues in rain-fed fig production areas is a lack of precipitation and drought. Most
fig trees in Iran are grown in Estahban, a semi-arid area in Fars province’s southeast.
The production of Iran dried figs is dominated by this area, which produces 90% of
figs (Javanmard & Mahmoudi, 2008; Javanmard, 2010). Extreme temperatures in
this region range between −7 and + 41 °C in winter and summer, respectively. The
annual rainfall averages around 350 mm years−1. The majority of the precipitation
falls in the late fall and winter (Rahemi & Jafari, 2005).
Brady and Weil (2002) found that more than half of the precipitation received in
semi-humid, semi-arid locations evaporates and returns to the atmosphere. In the
tropics, particularly in semiarid locations, water is frequently a key limiting factor
for agricultural production (Scopel et al., 2004). Furthermore, water shortage is the
most critical stress factor impacting plant development in most terrestrial ecosys-
tems (Kozlowski & Pallardy, 1996). Zare et al. (2009) performed socioeconomic
research in this region to investigate the effects of drought on rainfed fig farming.
The findings revealed that the damaged rain-fed orchards of Estahban had a 50%
loss in fig production and price, implying that the fig tree is directly or indirectly
responsible for 85% of the economy in the city. The quantity of precipitation in
Estahban is usually about 350 mm, and the amount of fig output is around 17,000 t.
In 2007, 64 mm annual rainfall in certain locations resulted in more than 80% crop
decrease. Due to a lack of water, around 10% of trees died this year, and the remain-
der had major physiological difficulties (Jafari et al., 2015).
270 M. Ben Temessek et al.

However, recent severe droughts have had a considerable impact on fig produc-
tivity. Furthermore, world water supplies are currently constrained because of
global climate change. Therefore, many researches have been applied to examine
the response of fig trees to drought stress and additional watering (Tapia et al., 2001;
Mi et al., 2009).
According to the authors, drought significantly reduces growth and productivity.
Furthermore, there had been a positive and significant association between irriga-
tion rates and vegetative growth or yield in all supplemental irrigation treatments
(Jafari et al., 2012). Researchers studied the edible fig ‘Sabz’ at the Estahban Fig
Research Station in Fars province from February 2008 to September 2011. The
effect of trunk thinning on drought diseases was studied using a random complete
block design with three replications.
Because ‘Sabz’ fig trees are trained with numerous trunks, 16 intensities of trunk
thinning were compared during winter pruning. The treatments were achieved by
trimming five separate trees to leave 0, 1, 2, 3, or 4 trunks tree−1, respectively. The
pruning intensity of each of the 16 treatments was calculated by dividing the total
circumference of the residual trunks by the total trunk circumference before the
thinning treatment was applied (Jafari et al., 2015). Different levels of trunk thin-
ning improved tree performance in terms of leaf width, leaf number, leaf water
potential, leaf temperature, leaf chlorosis, leaf necrosis, fruit color, and size.
The statistical analysis of the 2 years findings revealed that trunk thinning sub-
stantially impacted the quantity of leaves per tree. The number of leaves per plant in
0 T trees (trees with all trunks thinned) was much higher than in control trees and
other treatments. Trunk thinning had a considerable impact on leaf width (which
varied from 9.4 to 13.6 cm); the 0 T (one treatment contained trees with no trunks
remaining and all trunks removed from the soil surface using pruning) treatment
raised this parameter much more than the other treatments. Improvements in tree
water status caused by trunk trimming may have created a more favorable environ-
ment for these trees to produce bigger leaves. For growth to occur, turgor must be
maintained (Prajapati et al., 2003). Partial tree wilting can be caused by a reduction
in turgor. The loss of turgor, which affects the rate of cell growth and final cell size,
is the initial result of water stress. Loss of turgor pressure is probably the most sensi-
tive process to water stress. As a result, growth rate, stem elongation, leaf dilation,
and stomatal opening are reduced. It also significantly impacts yield quality
(Prajapati et al., 2003). They increased the root-to-shoot ratio by severe pruning
(trunk thinning); thus, leaf water potential increased.
Because of increased leaf transpiration, which can lower leaf temperature,
greater leaf water potential leads to reduced leaf sunburn (Mi et al., 2009). The more
leaves create a better cover and lower the temperature of the leaves under the can-
opy, so the sunburn on the leaves is also reduced. Although reducing the number of
trunks per tree under drought stress increased fig tree resistance to water stress,
thinning all the trunks enhanced the vegetative look more than the other treatments.
This treatment guarantees the survival of rain-fed fig trees in dry years. In general,
trunk thinning of rainfed fig trees improves vegetative and reproductive qualities by
reducing overall tree transpiration requirement and sink organs in drought
11 Fig (Ficus carica L.) Production and Yield 271

situations. This demonstrates that this kind of pruning (thinning the whole trunk)
had no positive influence on fruit growth in fruit yield and growth. Fruit size
decreases due to reduced cell size and division due to lower turgor pressure and
internal auxin content (Jones et al., 2008; Prajapati et al., 2003). Since the signifi-
cant increase in weight and size of edible figs occurs in the third phase of fruit
development, a favorable tree water status plays a major role during this phenologi-
cal stage (Mitra, 1997).
Tree water status plays a vital role in cultivating fig trees under rain-fed condi-
tions, improved by trunk thinning in rain-fed figs in dry years. Trunk thinning also
led to increased vegetative growth and fruit quality. In this study, thinning all of the
trunks produced the greatest advantages in terms of tree appearance. Furthermore,
it had a good impact on both tree and fruit qualities. As a result, producers working
in dry years are advised to use the later pruning method (Jafari et al., 2015).

10 Effect of In Vitro Thermotherapy Application on Shoot

Development in Some Fig Cultivars

The fig tree (Ficus carica L.) has piqued interest due to its economic and therapeutic
benefits. The fig tree has a great genetic variety in Tunisia, and all cultivars are well
adapted to local circumstances (Mars et al., 2008, 2009). However, several abiotic
and biotic restrictions hinder the growth of this fruit crop, contributing to a decline
in revenue and the progressive removal of plantations (Saddoud et al., 2011). The
scarcity of pollinators that have been specifically chosen to meet the demands of
female cultivars and the challenges in multiplying them in the field are severe
restrictions (Gaaliche et al., 2013). In addition, virus infections particularly fig
mosaic disease, pose a problem and have spread throughout the industry (Saddoud
et al., 2007), inhibiting the growth of healthy orchards and limiting production
(Ashihara et al., 2004). According to Gella et al. (1997), fig mosaic disease is fre-
quent in Mediterranean regions with temperate and subtropical climates. By defoli-
ating leaves and dropping fruit, the disease causes economic losses. Thermotherapy
and meristem culture are effective strategies for growing virus-free plants.
Thermotherapy can be used in one of three ways: (1) Thermotherapy is adminis-
tered to a plant first, and then meristem culture is formed; (2) Thermotherapy is
applied to in vitro shoots first, and then meristem/shoot tip culture is established; (3)
Thermotherapy is applied to cultured meristem/shoot tip (George, 1993). In vitro
thermotherapy on fig shoot cultures infected with fig, mosaic has demonstrated that
fig shoots produced in vitro are heat tolerant (Gella et al., 1997).
As well as that, Günver et al. (2015) used a thermal approach to treat the tips of
fig shoots, and they assessed the shoot viability ratio and the condition of shoot
development. So, 40 shoot tips (0.7–1.0 cm) of fig cultivars ‘Sarlop’, ‘Bursa Siyah’
female and ‘Ak ilek’, ‘Kaba ilek’ male was by hardwood cuttings with mosaic
symptoms. The explants were grown in MS (Murashige & Skoog, 1962) medium,
which included 2.0 mg L−1 benzyladenine (BA), and 0.2 mg L−1 gibberellic acid
272 M. Ben Temessek et al.

Table 11.4 Shoot ratio and new shoot emergence ratio (%) after thermotherapy applications
Survival ratio (%) New shoot emergence ratio (%)
Treatment Sarılop Bursa Siyahı Sarılop Bursa Siyahı
Control 67.00 69.69 50.00 46.00
Thermotherapy 35.00 55.00 10.00 23.00
Günver et al. (2015)

(GA3), 0.1 mg L−1 indole butyric acid (IBA), and 89 mg L−1 phloroglucinol (PG). As
a result, first, due to severe contamination, shoot explants of the male fig cultivars
‘Ak ilek,’ and ‘Kaba ilek’ were eliminated from the experiment.
Table 11.4 shows the shoot survival ratios of the female fig cultivars ‘Sarlop’ and
‘Bursa Siyah’ 3 weeks after thermotherapy treatments. After 5 eeks of thermother-
apy administration, the control had the highest survival ratio and the fresh shoot
emergence ratio excised from living shoot tips. Thus, the control group had a higher
shoot emergence ratio than the thermotherapy group. Bursa Siyah outperformed
Sarlop in thermotherapy applications among cultivars. Finally, when thermotherapy
was applied, the shoot survival ratio and new shoot emergence ratio were reduced
compared to the control. These ratios varied depending on the cultivar. On the other
hand, Gella et al. (1997) acquired a very high survival rate in fig shoot cultures the
following thermotherapy. The benefit of in vitro thermotherapy is that it produces
larger explants (>1 mm) than meristem culture.

11 GIS-Based Soil Suitability Evaluation for Fig Orchards

in Southern Peloponnese, Greece

Most fig orchards in the Southern Peloponnese are located on various soil types, and
fig production is generally regarded as a low input in a dry-farmed system. However,
as production costs continue to rise and harvesting accounts for more than half of
total production costs, farmers must adopt more efficient and cost-effective cultiva-
tion and management practices (Melgarejo et al., 2007). According to the study of
Melgarejo et al. (2007), fig cultivation is most often associated with a low profile
and marginal lands (Because fresh figs have a concise shelf life and are fragile, the
majority of the world’s fig production is dried or otherwise processed (Stover et al.,
2007). However, researchers and growers should pay more attention to this fruit‘s
excellent nutritional worth.
In recent decades, tree-planting systems have seen a trend toward reducing till-
age or maintaining vegetation cover to reduce soil erosion and degradation and
improve water retention (Landa et al., 2014). However, the soil’s physicochemical
properties and fertility parameters must be considered to apply this soil manage-
ment practice. Therefore, soil samples were collected from 45 fig orchards in the
Messinia and Laconia regions of the Southern Peloponnese. Soil samples were
dried to constant weight using forced air at ambient temperature < 36 °C, then
11 Fig (Ficus carica L.) Production and Yield 273

passed through a 2 mm sieve (fine soil). Nine soil physicochemical parameters were
measured for individual GIS-based map references: pH, saturation percent (SP),
particle size analysis (PSa), organic matter (OM), cation exchange capacity (CEC),
calcium (Ca), Magnesium (Mg), Potassium (K) and Phosphorus (P).
The results of the nine soil parameter studies provided information on the suit-
ability of the studied areas for fig cultivation. Only for the soil parameters that show
inhomogeneity among the studied fig orchards are tabulated data or maps provided
(Kotsiras et al., 2017).
In terms of pH suitability, the findings show that the majority (73.3%) of the 45
orchards fall into category 4 (suitable), and 17.8% fall into category 5 (highly suit-
able). This indicates that the pH levels of the majority of the soils studied (91.1%)
are suitable for this crop (Kotsiras et al., 2017). The cation exchange capacity of soil
is an essential indicator of its productivity and can be used to make nutrition recom-
mendations for phosphorus, potassium, and magnesium levels in soils of various
textures (Adams et al., 1994). According to this study’s CEC findings, 75.5% of fig
orchards are moderate to very appropriate (Kotsiras et al., 2017).
Compared to other fruits, figs are considered extremely high in calcium (Rao
et al., 2002). When examining the individual values of the fig orchards, it is clear
that a large percentage of the soils (43%) have Ca concentration levels greater than
4000 ppm, which is consistent with the finding that a large percentage of the studied
soils (approximately 60%) have a pH value between 7.6 and 8.0 (Kotsiras et al.,
2017). On the other hand, most fig orchards (51.1%) have low amounts of exchange-
able Mg, and only 17.8% of the soil had appropriate and highly suitable Mg levels
(Fig. 11.1).

12 Effect of Organic and Inorganic Fertilizers on Fruit

Characters, Quality, and Economics of Fig Production

The fig (Ficus carica) is a nutritional fruit high in fiber, potassium, calcium, and
iron, with values higher than bananas, grapes, oranges, strawberries, and apples
(Vinson, 1999). Therefore, a field experiment was carried out to investigate the
effect of organic and inorganic fertilizers on the fruit characteristics, quality, and
yield of fig. Kurubar et al. (2015) used soil with a pH of 8.29 and 0.44% organic
carbon, 265 kg ha−1 of available N, 38 kg ha−1 of available P2O5, and 325 kg ha−1 of
available K2O in their experiment. In addition, the control inorganic fertilizer (RDF)
containing NPK was compared to the other twelve treatments based on different
combinations and doses of farmyard manure (FYM), poultry manure, vermicom-
post, and RDF (Kurubar et al., 2017).
The application of organic manure and chemical fertilizers through T9 and T11
resulted in significantly higher fruit length (5.82 cm), diameter (5.57 cm), number
of fruits plant−1 (430.6), fruit yield plant−1 (19.4 kg), and ha−1 (12.97 t), as well as
improved quality in the form of titratable acidity (0.24%) and TSS: acid ratio
(78.5%) (Kurubar et al., 2017).
274 M. Ben Temessek et al.

Fig. 11.1 Exchangeable Mg mapping in Messinia and Laconia, respectively. On the maps, suit-
ability is indicated by a five-color coded key ranging from unsuitable (red) to highly acceptable
(dark green). (Kotsiras et al., 2017)

According to the results of the Kurubar and Allolli (2015), the crop‘s economic
concept revealed that both treatments (T9 and T11) were lucrative, with the maxi-
mum net profits of Rs.198207 (T9) and Rs.185102 (T11), respectively, reflecting a
higher Benefit: Cost (B: C) ratio of Rs.6.58 and Rs.6.26 (Table 11.5). The use of
RDF alone resulted in increased yield and quality of fruits; however, it is also clear
that RDF used in combination and smaller quantities results in better size, quality,
and quantity of fruits, as well as higher profit with increased yield by the use of T9
(Kurubar et al., 2017).
Various workers have reported the beneficial effects of farm yard manure and
poultry manure, either alone or in combination with inorganic fertilizers, in various
crops such as guava, acid lime, coconut, banana, papaya, and guava (Corrales
Garriga et al., 2000; Dubey & Yadava, 2001; Ray et al., 2008).

13 Shelf-Life Extension Using Modified Atmosphere

Packaging and Moisture Absorption

Due to their rapid ripening and great susceptibility to rot, fig has a short shelf life
after harvest. As a result, postharvest strategies that allow for the extension of post-
harvest life in fresh figs, such as modified atmosphere packing, are required (MAP)
(Colelli et al., 1991). In addition, moisture absorbers for relative humidity (RH)
control could be a viable strategy for maintaining fruit quality and safety by delay-
ing fruit rotting and softening.
Shirazi and Cameron (1992) found that 10 g of sorbitol, xylitol, NaCl, KCl, and
CaCl2 allowed ripe green tomato fruit to be kept at 20 °C while maintaining a con-
sistent relative humidity. The results obtained by Villalobos and Ansah (2015) show
that no significant difference was identified between fruits stored under MAP (gas
concentrations of 20 kPa CO2 and 20 kPa O2) and fruits stored in MAP with 3%
(w/w) of CaSO4, indicating that the use of this moisture absorber did not appear to
affect the fruit respiration rate. Other authors (Villaescusa & Gil, 2003) reported
similar results, stating that using moisture absorbers such as sorbitol or silica gel in
11 Fig (Ficus carica L.) Production and Yield 275

Table 11.5 Effect of organic manures and inorganic fertilizers on the economics of fig cultivation
Cost of cultivation (Rs. Gross returns (Rs. Net returns (Rs. B:C
Treatment ha−1) ha−1) ha−1) ratio
T1 37,096 195,030 157,934 5.26
T2 35,259 152,100 116,841 4.31
T3 36,487 171,090 134,603 4.69
T4 34,650 128,970 94,320 3.72
T5 33,950 176,850 142,900 5.21
T6 32,113 153,810 121,697 4.79
T7 36,792 210,240 173,448 5.71
T8 34,955 145,080 110,125 4.15
T9 35,523 233,730 198,207 6.58
T
10 33,686 179,820 146,134 5.34
T
11 35,218 220,320 185,102 6.26
T
12 33,380 133,020 99,640 3.98
T
13 33,248 167,130 133,882 5.03
T1: FYM at 16.5 t ha−1 + 75% RDF; T2: FYM at 16.5 t ha−1 + 50% RDF; T3: Vermicompost at
5 t ha−1 + 75% RDF; T4: Vermicompost at 5 t ha−1 + 50% RDF; T5: Poultry manure at 5 t ha−1 + 75%
RDF; T6: Poultry manure at 5 t ha−1 + 50% RDF; T7: FYM at 8.25 t ha−1 + vermicompost at
2.5 t ha−1 + 75% RDF; T8: FYM at 8.25 t ha−1 + vermicompost at 2.5 t ha−1 + 50% RDF; T9: FYM
at 8.25 t ha−1 + poultry manure at 2.5 t ha−1 + 75% RDF; T10: FYM at 8.25 t ha−1 + poultry manure
at 2.5 t ha−1 + 50% RDF; T11: Vermicompost at 2.5 t ha−1 + poultry manure at 2.5 t ha−1 + 75%
RDF; T12: Vermicompost at 2.5 t ha−1 + poultry manure at 2.5 t ha−1 + 50% RDF; T13: RDF alone
(200:132:132 NPK kg ha−1)

conjunction with MAP did not affect the headspace gas concentration. Del carmen
et al. (2014) reported minor weight losses in figs under MAP due to the film’s effect
on increasing vapor pressure. Furthermore, after 14 days of cold storage, the addi-
tion of CaSO4 showed no significant interactions with the type of film used. The
most significant weight loss was observed when fruits were stored with a humidity
absorb combined with a micro-perforated film.
High levels of CO2 and low levels of O2 have been shown to suppress mould and
yeast growth in fruit, particularly figs (Jacxsens et al., 2001; Crisosto & Kader 2007;
Bouzo et al., 2012). Furthermore, moisture may have antibacterial properties since
the amount of water available on the skin surface is likely to be lower than the essen-
tial water activity of microorganisms, which is an efficient approach to inhibiting
their growth (Falconi, 2008). Shirazi and Cameron (1992) reported similar results,
claiming that using NaCl as a moisture absorber in red ripe tomato fruit prevented
surface mold growth.
The use of MAP has been proven to be beneficial in maintaining the quality of
fig fruit and, as a result, extending its potential storability, reducing weight and
acceptability losses, and regulating microbial growth. However, even though CaSO4
application controlled fungal growth and resulted in the highest firmness, it nega-
tively influenced fruit attractiveness. As a result, no synergistic effects were detected
with moisture absorbers, which had minimal benefits when used without MAP
(Villalobos & Ansah, 2015).
276 M. Ben Temessek et al.

14 Developing the Fresh Fig Industry

Fig fruit, one of the world’s first cultivated trees, is grown in most temperate cli-
mates. Figs are sodium-free, fat-free, and cholesterol-free, just like other fruits. Figs
are also high in vitamins, amino acids, phenolics, and antioxidants (Solomon et al.,
2006). Unfortunately, fresh figs are extremely vulnerable to physical damage and
post-recollection infections, which cause significant losses during commercializa-
tion. Therefore, the conditions of verging before and after harvesting are critical for
improving fruit quality and shelf life after harvesting (Preece et al., 2015). An in-
store consumer review of four California fresh fig cultivars at two maturity stages
found that cultivar and maturity at harvest were the most critical factors affecting
like (acceptance).
Figs picked at tree-ripe maturity received a considerably higher acceptance rat-
ing than figs harvested at commercial maturity in all four cultivars. The average
acceptance rate for the four cultivars was 86% for tree-ripened figs and 55% for
commercially ripened figs (Crisosto et al., 2010). Therefore, it is time to find new
cultivars that can be harvested at an advanced or commercial maturity stage and
withstand postharvest processing. Unfortunately, there is a scarcity of high-quality
consumer cultivars that can be picked at an advanced maturity stage and remain
solid throughout postharvest handling. As a result, we looked for genotypes and
cultivars with low decay susceptibility and desirable sensory traits such as sweet-
ness, pleasant volatiles, and firm texture during postharvest handling (King et al.,
2012; Haug et al., 2013).
To promote fresh fig consumption in the United States and abroad, it is necessary
to choose high consumer-grade figs with good flavor that remain firm during ripen-
ing and survive postharvest handling (Crisosto et al., 2017). Therefore, the National
Clonal Germplasm Repository (NCGR) advised a breeding program for generating
firm-fleshed cultivars with high consumer approval based on postharvest analyses
of consumer acceptance, firmness, and decay susceptibility of promising fresh figs
(Preece et al., 2015).

15 Fresh, Semi-Dried and Sun-Dried Fig (Ficus carica

‘Sarılop’) Fruit Production

Due to the diversity of ecological conditions and geographical structures in Turkey,

fig cultivation is practiced in various regions of the country. It is an economically
important fruit species in Turkey’s Western Aegean region, particularly in the Great
and Lesser Meander Valleys (Arpaci, 2015).
The average fruit weight, the ratio of defective fruit (sunscald, cracked, or black
neck), and total soluble solids and glucose levels in fresh, semi-dried, and dried fig
fruit were significantly affected by climatic circumstances during fruit growth, mat-
uration, and drying. Furthermore, skin color C* and h° values in fresh figs; skin h°
11 Fig (Ficus carica L.) Production and Yield 277

value and total sugar contents in semi-dried figs; and firmness and total sugar con-
tents in dried figs were affected positively or negatively by climatic conditions.
The best results were obtained in 2011 when temperatures did not surpass 40 °C,
and relative humidity was between 40 and 70%. Between the beginning of August
and the end of September, the temperature/relative humidity regime significantly
impacts overall dried fig quantity and quality in the Region, particularly fruit weight
and defect ratio. Some of the unfavorable consequences, such as high relative
humidity levels or rain during the sun-drying period, can be partially mitigated by
using highly aerated tunnels for drying. Sun drying is a cost-effective and ecologi-
cally friendly preservation method, and modest management changes can improve
dried fig fruit‘s ultimate quality and marketability (Şen et al., 2017).

16 Conclusion

The global production of the fig (Ficus carica L.) has grown in prominence, owing
to its significant function as a dietary supplement. This circumstance has sparked
much curiosity about its manufacturing, processing, and preservation methods.
Growing fig trees in poor circumstances can result in crop loss and numerous fruit
damage. Although this crop may grow in various climates, agronomic consider-
ations such as production method, fertilizer management, water demand, and soil
requirement can all impact fig tree yield and output. Increasing the value of agricul-
tural products cultivated, inefficient production systems are required to provide a
good return on investment. Thus, proper agronomic management must be employed
to eliminate difficulties caused by internal and external causes, creating an environ-
ment suited for operating efficiency and improved Ficus carica L. output.

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Chapter 12
Defense Mechanism of Fig (Ficus carica)
Against Biotic Stresses: An Advanced Role
Model Under Moraceae

Sudeepta Pattanayak, Siddhartha Das, and Suryakant Manik

Abbreviations

ISR Induced Systemic Resistance

PAG Psoralic acid glucoside
ROS Reactive oxygen species
SAR Systemic Acquired Resistance

1 Introduction

The genus Ficus is one of the widely versatile members, which comprises approxi-
mately 800 species of herb, shrub, trees, and epiphytes and are distributed cosmo-
political (Singh et al., 2011). Ficus carica (Moraceae) is a large deciduous member
belonging to Moraceae with a unique defense system. Ficus has wide adaptability
through its various lifeforms like epiphytes, trees, and vines in various climatic and
geographic regions (Pierantoni et al., 2018). Morphological uniqueness is visible
with its unique inflorescence type (syconium). The most well-known Ficus species
is Ficus carica, which is deciduous and maintains a height of up to 5 m. The origin
Ficus thinks to be Middle East countries which latterly distributed in warmer
regions across the globe. A maximum diverse population exists on the Asian and
Australian continents. Different names of Ficus in different locations are reported
(Table 12.1). In alternate Ficus population in Africa is less, approximately 100–150
(Badgujar et al., 2014). Fruits of Ficus are edible and could be consumed in fresh,

S. Pattanayak · S. Manik
Division of Plant Pathology, ICAR-Indian Agricultural Research Institute, New Delhi, India
S. Das (*)
Department of Plant Pathology, M.S. Swaminathan School of Agriculture, Centurion
University of Technology and Management, Paralakhemundi, Odisha, India

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 283
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_12
284 S. Pattanayak et al.

Table 12.1 Various names of Ficus carica across the globe

Sl. No Country/location Name
1. India Fig, Dumur, Anjir, Angir, Shimayatti
2 China Mo Fa Guo, Wu Hua Guo
3. Czech Smokvon
4. German Echter, Feigenbaum, Essfeige
5. Italian Fico, Fico Comune
6. Iran Anjeer
7. Korea Mu Hwa Gwa, Mu Hwa Gwa Na Mu
8. Russia Inzir
9. Sweden Fikon, Fikontrad
10. Turkey Incir, Yemis

dried, and processed forms by all classes of people. Although, different countries
like Turkey, Egypt, and Algeria are the pioneer in promoting Ficus cultivation
commercially.
Ficus carica, therefore, is considered an important species with both agronomi-
cal and economic impacts. Several Ficus species showed wide genetical diversity
with excellent pharmacological potentiality. Like other plant species, Ficus is also
infected by several destructive insect pests (Novtny et al., 2010), fungi (Hosomi
et al., 2012), and viruses (Bayoudh et al., 2017). Though a few numbers of patho-
genic microbes infect Ficus severity or aggressiveness of the pathogen turned to
blur by its unique defense mechanism (Chamont, 2014). Phytochemical studies
revealed that it is a rich source of phenolics, organic acids, and various volatile
compounds (Mawa et al., 2013). In the red cultivar of figs higher level of anthocy-
anin was observed. Carotenoids are mainly stored on the fruit peel. Ficus species
have developed various physical defense strategies like mineral-fiber and rigidity
levels. Chemical defense strategies encompass target-based metabolites and PR
proteins to combat foreign pathogens. Secretion of volatile organic compounds may
help Ficus racemosa attract parasitoids like Apocryptophagus agraensis and
Apocrypta westwoodi (Borges et al., 2013), which indirectly helps to control vari-
ous destructive insect pests. In this present chapter, we focus on a unique defense
system as a role model under Moraceae.

2 Physical Barriers or Structural Defenses

2.1 Hard, Tough and Mineralized Leaf Texture

Leaves are the herbivores’ main target, but various associated factors are directly or
indirectly responsible for the herbivores’ edibility property or food quality. Leaf
rigidity and abrasiveness are important defense indicators for a plant’s defense strat-
egy. Toughness creates a major barrier to herbivore crop consumption, while host
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 285

resistance might be related to storing biochemical substances such as lignin, fiber,

and phenolic content (Xiang & Chen, 2004). Toughness comes from the rebuilt
prolonged storage of calcium oxalate, silica, and calcium carbonate deposition. In
2018, Pierantoni and his co-workers reported that aforesaid three components were
impregnated in various orientations among different species of Ficus in various
environmental conditions.

2.2 Calcium Oxalate Crystal-Based Defense

Calcium oxalate is deposited as a solid mineral on the plant surface (Korth et al.,
2006; Bauer et al., 2011). Calcium oxalate presented intra-cellularly and extra-
cellularly within the plant cells and gathered and impregnated in the vacuole, known
as idioblasts (Franceschi and Nakata 2005; Bauer et al., 2011). Idioblasts cells are
structurally and functionally different from the neighbor cells. These idioblasts cells
can develop in any part of the host. They can participate in multifold activities like
the calcium regulatory pathway and metal detoxifying pathway, which have a
broader defense mechanism impact (Franceschi & Nakata, 2005; Pierantoni et al.,
2018). Those crystals can develop a protective shield (Ruiz et al., 2002; Hudgins
et al., 2003) against destructive pests, and herbivorous infestation (Ward et al.,
1997) and are an effective corrading agent for the mouthparts of suckers and chew-
ing insects (Korth et al., 2006). After all, these calcium oxalate crystals turn into a
tough barrier in the defense mechanism (Finley, 1999). In Ficus, a fruitful eradica-
tion of pest infestation has been reported through the aforesaid defensive mecha-
nisms (Sosnovsky, 2016).
Calcium oxalate crystals are impregnated in parallel lines nearer to the main
veins of Ficus leaves. Hence the crystals are associated with the vein bundles, the
phloem, or the basal tissue of the veins and veinlets (Pierantoni et al., 2018). As per
the existence in different locations within the plant body, those calcium oxalate
crystals are well-known as prismatic crystals, extended to mesophyll and bundle
sheath cells (Wu & Kuo-Huang, 1997; Pierantoni et al., 2018). Occurrence of cal-
cium oxalate crystal founds in bundle sheath cell and mesophyll cells in the form of
druses in Ficus variagata. In the case of Ficus elastica, the entire mesophyll layer
and bundle sheath surface are covered with the sharper needle-like calcium oxalate
crystals, which are also dispersed throughout the epidermal region of leaves (Wu &
Kuo-Huang, 1997). In F. carica, calcium oxalate was noticed in the phloem region
in two different shapes – druses and prismatic crystals. Developmental and deposi-
tional orientation patterns revealed that calcium oxalate was stored on both dorsi-
ventral sides of the leaf vein and covered the entire surface (Pierantoni et al., 2018).
In Ficus species, calcium oxalate crystals exist as prismatic crystals for their local-
ization towards bundle sheath. However, similar several other variations are also
observed. In the case of Ficus elastica, two different types of crystals are found;
prismatic crystals are larger numbers towards bundle sheath cells, whereas
286 S. Pattanayak et al.

needle-shaped in fewer numbers (Wu & Kuo-Huang, 1997). Occurrence of calcium

oxalate crystal found in F. carica in the phloem. Calcium oxalate crystals are depos-
ited on the dorsi-ventral surface of the leaf.

2.3 Calcium Carbonate (Cystoliths) Based Structural Defense

Calcium carbonate is one of the major components which originates and provides
structural defense in some specific members of Moraceae. These calcium-oriented
depositions made up a unique layer of cellulose microfibrils with pectin. Cystoliths
are oblong, reticulate, cigar, ‘Y- shaped’, I shaped, bean-shaped, grapes-shaped, etc.
The size of the cystolith crystal varies from 20–80 μm in wide and 10–120 μm in
length. The shape of cystolith depends on its localized occurrence in a specific part
of adaxial tissue or abaxial tissue. In the case of Ficus benghalensis, cystolith is
found in adaxial condition, and Ficus religiosa is in abaxial condition (Pierantoni
et al., 2018). Based on that, it transformed into an elongated or spherical cystolith.
Mainly, the occurrence of Cystolith is noticed in idioblastic cells and is well known
as lithocysts. Lithocysts frequently showed an externally projected sharper tail
made up of silica whose size depends on silica deposition on the surface.

2.4 Silica Compositions

Silicon is easily available under the soil and easily intake by plants (Strömberg
et al., 2016). Silicon helps in multifold activities like in various physiological pro-
cesses and makes a protection shield against various biotic stresses. Various polym-
erized silica in plant bodies was the toughest element that helped resist plants.

2.5 Crystal Enriched Idioblast Cell

In Ficus carica leaves, several idioblasts that comprise spiny crystals were found
along with the primary layer of palisade cells. Those crystals are unique structures
in the cystoliths, but characterization has not been diagnosed (Mamoucha et al.,
2016). However, those sharper crystals would possibly create leaf acerbity and
harshness.
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 287

3 Specialized Transformed Structures

3.1 Glandular Trichomes

Glandular trichomes are multi-celled structures that project externally from the
main plant surface, secreting viscous substances of defense-related metabolites.
Morphological variability is diverse among the Ficus species. However, character-
istic features of glandular trichomes are uniseriate, capitates, and filamentous.
Capitate types of trichomes bear a short stalk (nearly 5–6 celled). In the case of
Ficus erecta, glandular trichomes are elliptic, and in Ficus sagitata, it is spherical.
In Ficus, glandular trichomes are present in both dorsi-ventral surfaces, but the
maximum occurrence is found on the ventral surface (Sosnovsky, 2016). In the case
of Ficus carica, glandular trichomes are very small, approximately 4 celled struc-
tures. The origin of these small appendages occurred at the time of maturity of
the leaves.

3.2 Non-glandular Trichomes

Trichomes are generally two types; glandular types and non-glandular types. Non-
glandular trichomes are one of the essential structures and cosmopolitical present in
almost all the Moraceae members, which helps in structural defense mechanisms.
These celled structures are also well-known as idioblasts. Non-glandular trichomes
make one structural shield against herbivores and other similar agents. Within the
Ficus genus, initially growing lengthy trichomes seem to be major and extra ample
on the abaxial facet than on the adaxial side of the leaf; however, their precise shape,
length, and density were found to be stronger with inter- and intra-specific varia-
tions (Klimko & Truchan, 2006). Small unicellular hairs containing silica deposi-
tion covered the leaf‘s surfaces, although maximum growth is found on the adaxial
surface. Some peltate non-glandular trichomes are found on the abaxial surface. In
F. carica leaves, minute non-glandular trichomes appear at initial developmental
stages and alter their external appearance by competitive look earlier than they pro-
gressively appear on the leaf surfaces.

3.3 Laticifers

Laticifers are a specialized secretory structure or a group of connected cells contain-

ing a liquid-like substance known as latex. These are a complex network of elon-
gated living cells spread through the veins in entire plants. The physical structure
and anatomy mainly vary from species to species. Similarly, the concentration of
latex secretion varies from low to high among species (Kim et al., 2003). Latex is a
288 S. Pattanayak et al.

major constituent of many potential proteins and enzymes that play a key role in the
plant-pathogen war. These laticifers are reported mainly from tropical plants having
a significant role in defense strategy (Hagel et al., 2008; Konno, 2011). For exam-
ple, in the plant Ficus, a cysteine protease (ficin) plays a major role in defense
against several pests (Kajii et al., 2014). Though laticifers can be articulated or non-
articulated (Fig. 12.1), in the genus Ficus, these are non-articulated and contain
more than one nucleus. These laticifer cells are mainly observed during the embry-
onic stage and confined to regions around the leaf veins only where the secretory
system of capitate trichome is also present. Therefore, it clarifies the presence of
these structures just above the xylem and near the nerves (Mamoucha et al., 2016).
Latex is a constituent of various chemicals that can be a potential weapon against
pests, pathogens, and herbivores; it can act as a physical barrier and chemical
defense simultaneously. Even very few amounts of latex are present in some Species
if the Ficus plant can effectively control or defend the pests and herbivores. For
instance, latex is present in laticifers cells under high pressure (Agarwal & Konno,
2009). When an injury occurs by any biotic agents, a sudden rupture of laticifer cells
occurs, resulting in the secretion of latex in a high amount (Farrell et al., 1990). The
sudden secretion of lates in a larger concentration can potentially control pests and
mammalian herbivores (Konno et al., 2004).
Moreover, when the latex comes in contact with the external environment or air,
it becomes dry and sticky, forming a physical defense barrier against a wide range
of pests. The stickiness and color of the latex are due to the presence of a terpenoid
or rubber, which is secreted from laticifer cells. This may be closing the injuries to
stop the entry of harmful biotic agents or immobilizing harmful insects by trapping
them or their other body parts (Dussourd & Denno, 1994; Kim et al., 2003).
Additionally, the presence of chemicals such as secondary metabolites, irritant
properties, enzymes, etc., in higher concentrations than other plant parts has made

Fig. 12.1 Structure and anatomy of articulated leaves (a) and non-articulated leaves (b). Ficus
plants have non-articulated laticifer
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 289

them more potential defense structure in the whole plant. The chemical defense of
latex has also been reported from Ficus carica (Kitajima et al., 2018) and Ficus
virgata (Konno et al., 2004). In the case of Ficus carica, more than 50 proteins have
been observed, and many of them are reported to have defensive proteins (Kitajima
et al., 2018). Similarly, in the Ficus virgata plant, the compounds present in latex
can effectively control lepidopteran larvae (Konno et al., 2004).

4 Cellular Defense Responses

During host-pathogen/pest interaction, the physical and chemical barrier of the host
serves as the first line of defense and protects itself from the harmful effect of such
introducing or interacting pests. However, in some cases, or due to a lack of the first
line of defense, one or both defensive barriers fail(s) to counteract such virulent or
aggressive pests. Then, the host plant develops an active and dynamic mechanism to
gain cellular resistance, which is termed cellular defense. Cellular defense responses
are categorized into two groups (Fig. 12.2) based on their time and duration of
occurrence: – (i) Rapid active cellular defense and (ii) Delayed active cellular
defense.

Fig. 12.2 Mechanism of host-pathogen interaction and development of rapid and delayed defence
responses in plants against the pathogen. [SAR (Systemic Acquired Resistance) and ISR (Induced
Systemic Resistance)]
290 S. Pattanayak et al.

4.1 Rapid Active Cellular Defense

Activation of rapid cellular defense resides in the consequences of host-pathogen

interaction. Understanding their molecular level interaction, there is a need to rec-
ognize elicitors (Pathogen’s avirulent gene product) and with the corresponding
receptors (host’s R gene product) and formation of receptor- elicitor complex, which
will trigger rapid response cellular defense. This complex causes quick and delayed
responses by triggering signal transduction reactions at the local and systemic lev-
els, respectively.

4.2 Delayed Active Cellular Defense

This defense system is activated in the host later to the activation of rapid cellular
defense, and it is systemic and persists for a more extended period. It occurs in
plants within minutes to hours of establishing the elicitor-receptor complex.
Events of Rapid Active Cellular Defense
Plants have an integrated signalling system that regulates plant growth and develop-
ment by mediating perceptions and responses to hormones, nutrients, the environ-
ment, and stressors. Plant-pathogen interactions trigger a range of plant defense
mechanisms (Hammond-Kosack & Jones, 1996), and these interactions are compli-
cated, and there is no general model or sequence of events that accurately represents
the dynamics of resistance. When the elicitor ligand binds to its receptor, it triggers
the cascade of signalling phenomena such as protein phosphorylation, ion fluxes,
activation of reactive oxygen species (ROS), nitric oxide (NO), etc. These signalling
pathways bring about different events of cellular defense, which include:

4.2.1 Alternation of Membrane Function

According to most studies on the initial stages of the host-parasite interaction, it was
demonstrated that the host membrane plays a vital role in pathogen identification
and signal transduction. Alteration in the permeability of plasma membrane led to
proton (H+) and calcium (Ca2+) influx as well as chloride (Cl−) and potassium (K+)
outflow in plant cells (McDowell & Dangl 2000). In addition, ion fluxes have a
significant role in activating reactive oxygen species during host-pathogen
interaction.
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 291

4.2.2 Oxidative Burst

Oxidative burst is significant defense machinery in host-pathogen conflict (Alvarez

et al., 1998). Extracellular synthesis of reactive oxygen intermediates such as super-
oxide (O2−), hydroxyl free radical (OH−), and hydrogen peroxide (H2O2), known as
oxidative burst. It is induced by ion fluxes and mediated by NADPH oxidase and/or
apoplastic-localized peroxidases (plasma membrane-located) (Somssich &
Hahlbrock, 1998; Abdin et al., 2013). Following pathogen attack, cell wall peroxi-
dases have also been implicated in producing reactive oxygen species (ROS)
(Bolwell et al., 1995). The fate of oxidative burst in plant cell include:
• Expression of defense genes
• Cross-linking of host cell wall
• Direct killing of invading pathogen at their site of infection
• Triggering of the hypersensitive response in plant cell
• Moderate to severe host cell damage

4.2.3 Reinforcement of Cell Wall

To establish a parasitic relationship with the host plant, most pathogens must breach
host cell walls at some stages, either as germ tubes, hyphae, or haustoria. To coun-
teract the ingress of the pathogen, there is an increase in cytoplasmic streaming and
accumulation of such cytoplasmic aggregates in infected cells of the host, which are
promised to strengthen the cell wall. Cell wall strengthening or fortification of the
cell wall is carried out by forming cell aggregates, including papilla, lignotubers
(lignified callose deposits), and hydroxyproline-rich glycoproteins. Reinforcements
of the cell wall, due to infection of the pathogen, in a resistant host plant are rapid,
and the structure formed is larger, where an incompatible host-pathogen interaction
takes place, and in the case of a susceptible host, this process is delayed, and struc-
ture formed is smaller in size providing a narrow range of protection (Guest &
Brown, 1997).

4.2.4 Hypersensitive Response (HR)

Hypersensitive Response is defined as the rapid death of cells in the immediate and
surrounding regions of an infection, which acts to limit the growth and development
of the pathogen and transmission to other regions of the plant. E.C. Stakman
(Stakman, 1915) is known to be the first person to coin the word “hypersensitive.”
He stated that the ‘host plant in such situations is hypersensitive to the fungus’ when
discussing the extreme resistance of particular grass hosts to Puccinia graminis.
This phenomenon of disease resistance development in plants is similar to apopto-
sis, or programmed cell death, which is observed during the development of disease
defense in mammals. Hypersensitive cell death typically occurs before oxidative
292 S. Pattanayak et al.

burst, increased cytoplasmic aggregation, streaming, granulation, membrane break-

down, and cellular decompartmentalization, all occurring within 12–24 h of
attempted entry (Guest & Brown, 1997). HR is a common occurrence in most, if not
all, higher plants, and a variety of pathogens can trigger it. HRs caused by a fungus,
oomycetes, bacteria, and viruses are the most prevalent, but HRs can also be gener-
ated by other species that create persistent intimate connections with the host plant,
such as insects (Rossi et al., 1998) and nematodes (Dropkin, 1969). For example, a
link between necrotic mesophyll cells in Bromus sp. and the fungus Puccinia recon-
dite was noted, attempting to infect resistance cultivars (Guest & Brown, 1997).

4.2.5 Phytoalexin Accumulation

One of the most significant advances in physiological plant pathology in the last
25 years has probably been the phytoalexin paradigm of disease resistance. Plants
produce phytoalexins, which are low-molecular-weight secondary metabolites that
serve as antibiotics in response to pathogen infection, and they are undetectable in
uninfected plant tissues. The accumulation of defensive chemicals is generally fol-
lowed by the hypersensitive response of infected host cells. This phytoalexin-
mediated resistance response of the host is considered the last event of rapid cellular
defense and subsequently activates a wide range of delayed resistance phenomena
in the host plant. More than 300 phytoalexin-like compounds have been identified
from plants belonging to over 30 different families. Phytoalexins produced by plants
in the same family usually have comparable chemical structures; for example, phy-
toalexins produced by most legumes are isoflavonoids, while phytoalexins produced
by the Solanaceae are terpenoids. The majority of phytoalexins are produced in
plants in response to fungus infection. However, phytoalexins have also been
induced by bacteria, viruses, and nematodes, and their mode of action includes: (i)
disruption of membrane integrity and function and (ii) Inhibition of the electron
transport system (Huang, 2001).
Ficus species are infected by fungal, viral, and bacterial pathogens, which cause
minor to major crop losses. The most common diseases of fig include Bacterial
canker, fig rust, anthracnose, and fig mosaic, and the occurrence and severity of
diseases vary with different places. It is reported that fig canker (by bacteria
Xanthomonas campestris) and fig rust (by fungus Cerotelium fici) are the most com-
mon diseases in Europe and come to insect pest attack severe damage to fig is
caused by fig wax scale. In order to acclimatize themselves to the adverse biotic
stresses, Ficus species have modified their physical structures (like tough and min-
eralized leaves, glandular and non-glandular trichome, and laticifer) and also chem-
ical compositions (alkaloids, terpenoids, and phenolics) and accumulation of
defensive proteins like chitinase, protease inhibitors and PR (Pathogenesis Related)
proteins (Villard et al., 2019). Such a wide range of defensive activities of Ficus
may be achieved by constitutively and or by induced defenses. Accumulation of
some potent shielding chemicals and proteins in Ficus is mainly followed by a rapid
induced defense strategy method and results in delayed systemic and persistent
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 293

induced defenses. So far, there are no reports of the specific molecular mechanisms
unraveling the host-pathogen and host insect relationship in Ficus, but the occur-
rence of this is the most needed phenomenon to bring about rapid active cellular
defenses to ward off the pest attack.

5 Formation of Defensive Metabolites and Proteins

as Biochemical Defense

The genus Ficus contains many specialized potential chemicals and secondary
metabolites that act against various microbes and nematodes (Table 12.2) (Jeong
et al., 2009; Liu et al., 2011). In addition, these metabolites also confer other bio-
logical roles, such as irritant and antipyretic properties (Saeed & Sabir, 2002; Patil
et al., 2010). The number of proteins and unique compounds like terpenoids, alka-
loids, phenolics, etc., varies depending on the plant parts, species, cultivars, and the
existing environmental factors.

5.1 The potential Metabolites

Past research has proved that more than 100 metabolites have a role in defense-
related activities (Sirisha et al., 2010; Marrelli et al., 2014), while others are associ-
ated with specific biological properties like antioxidant activities (Oliveira et al.,
2009, 2012). Furthermore, the metabolites responsible for defense-related actions
are specialized, such as alkaloids, phenolics, terpenoids, etc.

5.1.1 Alkaloids

Alkaloid metabolites present in the Ficus genus have a wide range of defense activi-
ties such as antifungal, antibacterial, antiviral, anti-amoebic, and against a wide
range of insect pests (Bhutani et al., 1987; Baumgartner et al., 1990; Li et al., 2006).
Among all the alkaloid metabolites, the most studied one is the phenanthroindoli-
zidines, mainly found in Moraceae and Asclepiadaceae families (Al-Khdhairawi
et al., 2017). However, some less explored also hold potential defensive roles against
fungi and bacteria (Yap et al., 2016). These molecules are mainly confined to stems
and roots, sometimes leaves of Ficus septica (Wu et al., 2002; Damu et al., 2009),
Ficus hispida (Yap et al., 2015), Ficus fistulosa (Peraza-Sanchez et al., 2002; Yap
et al., 2016). It is also observed that the concentration of phenanthroindolizidines in
the latex and other plant exudates of some of the Ficus spp. much higher than it can
quickly kill herbivores (Konno, 2011). These compounds also display a strong cyto-
toxic effect on cancer cells (Peraza-Sanchez et al., 2002; Yap et al., 2016).
294

Table 12.2 Species of Ficus plants and their associated secondary metabolites with a defensive role
Secondary
metabolites or
Plant compounds Type Sub type Defensive Function References
Ficus Alkaloids Phenanthroindolizidine alkaloids Antofine Antifungal and Damu et al. (2009)
septica antibacterial
Ficuseptines A and B, 13aR- Toxicity toward cells Wu et al. (2002) and
tylocrebrine N-oxide (Cytotoxicity) Damu et al. (2005)
Indolizidine alkaloids 4,6-bis-(4-Methoxyphenyl)-1,2,3- Anti-bacterial and Baumgartner et al. (1990)
trihydrolndolizidinium chloride anti-fungal
Ficus Triterpenoids Methyl maslinate, oleanolic acid Irritant Jaina et al., (2013)
carica Steroids 6-O-acyl-β-d-Glucosyl-β-sitosterol Toxicity toward cells Rubnov et al. (2001)
(Cytotoxicity)
Monoterpenes, sesquiterpenes – – Ahmad et al. (2011) and
and esters, Ojo et al. (2014)
Phenolics Phenolic acids Chlorogenic acid Antioxidant Lazreg Aref et al. (2011)
Flavonoids Flavones, Flavanones, Rutin Antioxidant Vaya and Mahmood
anthocyanidins (2006) and Takahashi
et al. (2014)
Furanocoumarins Bergapten Nematicidal Alam et al. (2015) and
Guo et al. (2016)
Bergaptol, Xanthotoxin, – Mamoucha et al. (2016)
Xnthotoxol Chalepin, and Wang et al. (2017)
Ficus Alkaloids Chlorophenanthroindolizidine Tengechlorenine Toxicity toward cells Al-Khdhairawi et al.
fistulosa alkaloids (Cytotoxicity) (2017)
Septicine-type alkaloids Fistulopsine A and B Toxicity toward cells Yap et al. (2016)
(Cytotoxicity)
Ficus Flavonoids Pyranocoumarins 3-Hydroxyxanthyletin Antimycobacterial Chen et al. (2010)
nervosa
S. Pattanayak et al.
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 295

5.1.2 Terpenoids

The potential terpenoids such as norisoprenoids, esters, steroids, monoterpenoids,

triterpenoids, and sesquiterpenoids are present in most Ficus spp. These terpenoids
have been studied and observed to carry potential bioactive roles. Among all the
terpenoid compounds, the most studied one is sesquiterpenes and monoterpenes,
which are abundantly present in the leaves and fruits of Ficus carica, respectively.
The most significant example of defensive terpenoids in Ficus is cis-1,4-isoprene
polymer, popularly known as rubber, responsible for latex stickiness (Oliveira et al.,
2010; Konno, 2011).
Triterpenoids have also shown their promising roles in host defense. The three
most studied triterpenoids in Ficus spp. are methyl maslinate, lupeol acetate, and
calotropenyl acetate, which are present in the leaves and exhibit irritant activities
(Saeed & Sabir, 2002).
Similarly, antiproliferative roles were found in Ficus spp. due to the presence of
potential steroids. Many researches have been carried out to check the potentiality
of terpenoids. It was found that cardiac steroids, i.e., cardenolides are present in the
leaves of Ficus asperifolia (Roberts et al., 2016) while 6-O-acyl-β-d-glucosyl-β-
sitosterol is present in the latex of Ficus carica (Ojo et al., 2014; Rubnov et al.,
2001). Cardenolides show high toxicity against animals (Roberts et al., 2016) and
caterpillars (Konno, 2011). Cardenolides can cause several symptoms in caterpil-
lars, such as unresponsiveness, regurgitation, and convulsions (Dussourd & Hoyle,
2000). Another exciting fact observed by scientists is that many potential volatile
terpenoid compounds are also capable of showing significant defensive activity in
other plants. For example, antibacterial and anti-insecticidal activity were exerted
by monoterpenoids limonene which is present in the fruits of Ficus carica (Hebeish
et al., 2008). Other monoterpenoids present in the fruit are β-caryophyllene and
menthol, which can also act against bacteria (Kim et al., 2008; Skalicka-Wozniak
et al., 2009).

5.1.3 Phenolics

Phenolic compounds are mainly uniformly distributed in fruits and carry specific
properties like color, flavor, and defense against biotic stresses (Badgujar et al.,
2014; Villard et al., 2019). Ficus genus’s most studied phenolic compounds are tan-
nins, flavonoids, phenolic acids, coumarins, and derivatives. These are also known
for their antioxidant (Marrelli et al., 2014), antibacterial (Debib et al., 2014), and
anti-mycobacterial activity (Chen et al., 2010). For instance, flavonoids class of
compounds like genistein and pyranocoumarins such as 3-hydroxyxanthyletin pres-
ent in the root region of Ficus nervosa are known to have anti-mycobacterial activ-
ity (Chen et al., 2010). Similarly, the antibacterial activity of some of the fig extracts
may be due to the presence of flavonoids and tannins (Debib et al., 2014). Moreover,
one of the coumarins derivative, furanocoumarins, plays a potential defensive
296 S. Pattanayak et al.

compound against various plants (Ojala et al., 2000), herbivores, and pathogens
(Berenbaum, 2002; Guo et al., 2016).
The two significant furanocoumarins, such as Bergapten and Psoralen, are
reported from the leaves of Ficus spp. (Oliveira et al., 2012). This compound is
mainly located surrounding the nerves having glandular trichomes and laticifer cells
(Wang et al., 2017; Mamoucha et al., 2016). They could activate toxic reactive oxy-
gen species and cause mutation or cell death through a series of events (Dardalhon
et al., 1998; Berenbaum, 2002). Additionally, the compounds Bergapten and
Psoralen in figs plant can inhibit digestive enzymes such as amylase and cellulase,
resulting in a nematicidal effect (Guo et al., 2016). The photosensitizing action of
fig plants can also cause allergy-like contact dermatitis (Pathak et al., 1962). The
recent discovery of the presence of psoralic acid glucoside (PAG) in large concen-
trations many opened the path to find its physiological function (Takahashi et al.,
2017) and defensive role (Wang et al., 2017).

5.2 Defensive Proteins Present in Ficus species

The proteins such as chitinase, oxidase, protease, and protease inhibitors are present
in the Ficus plant lates and are very stable. Moreover, their properties are indepen-
dent and not affected by herbivores. Therefore, these proteins play a significant role
against pathogens and herbivores (Zhu-Salzman et al., 2008; Konno, 2011).

5.2.1 Chitinases

Chitinase is widely known to inhibit the growth of fungi by degrading the cell wall
(Taira et al., 2005). These can also show insecticidal activities sometimes by slow-
ing down their developmental process (Lawrence & Novak, 2006). Chitinase is
mainly found in the latex of Ficus microcarpa and Ficus carica (Kitajima
et al., 2018).

5.2.2 Oxidases

The main oxidase proteins, such as polyphenol oxidases and peroxidases, are pres-
ent in the Ficus genus, and they are associated with rubber and the rate of latex
coagulation, respectively (Wahler et al., 2009). The protein Peroxidases are mainly
found in the latex region of Ficus sycomorus, Ficus carica, and Ficus glabrata,
while polyphenol oxidases have been found only in Ficus elastica (Kim et al., 2008;
Mohamed et al., 2011). These proteins are considered one of the potential defensive
proteins due to their anti-insecticidal and antinutritive effects (Zhu-Salzman et al.,
2008; Konno, 2011). However, insects consume the Ficus plant parts, and a cascade
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 297

of signaling events occurs. This involves the production of ROS (reactive oxygen
species) for disruption of major amino acids and nutrients for the insects (Sethi
et al., 2009).

5.2.3 Proteases

Plant genomes encode some proteases whose key role is in host defense against
various harmful biotic and abiotic stress agents through generating signaling mole-
cules. For example, ficins, one cysteine class of protease, have been reported from
the latex of the Ficus family (Sgarbieri et al., 1964; Villard et al., 2019). This can
control many insects by disturbing their feeding habit, digestion, growth rate, and
development (Konno et al., 2004). For example, ficins in Ficus virgata can effec-
tively control lepidopteran insects in the larval stage (Konno et al., 2004).

5.2.4 Protease Inhibitors

Many plant species produce protease inhibitors, a major class of defensive protein
that is activated upon sensing a wound response initiated by an insect, pathogen, or
nematode. These are highly potential in defending the biotic agents, especially
restricting the infection by nematodes (Koiwa et al., 1997; Hellinger & Gruber,
2019). The major protease inhibitors are cysteine and serine protease inhibitors act-
ing as an important weapon against pests and pathogens (Kitajima et al., 2018;
Villard et al., 2019). Several evidences suggest the potential defensive role of tryp-
sin, a serine protease against insects and fungi (Ryan, 1990; Terras et al., 1993). The
latex of the Ficus carica plant contains different protease inhibitors, but the major
one is trypsin inhibitors, whose concentration is more and can target the proteases
present in insect gut (Huynh et al., 1992). Along with ficins, more than 10 isoforms
of trypsin inhibitors can be found in the latex of leaves.

5.3 Some Important Pathogenesis-Related Proteins (PRs)

Pathogenesis-related proteins, or PR proteins, have a significant role in the host

defense mechanism and are produced in host plants during various events of a
pathogen attack. The function and mechanism of these proteins are widely studied.
Significant PR proteins have a defensive role against fungi, bacteria, virus, and
insects (Ali et al., 2018). The major PR proteins found in Ficus plants belong to
PR-1, PR-4, and PR-5, and these are present mostly on the leaf latex (de Freitas
et al., 2011). In addition, these proteins can inhibit or restrict the growth of many
pathogens, mostly fungi (Bai et al., 2013), hence considered fungicides.
298 S. Pattanayak et al.

Table 12.3 Chemicals and proteins present in the latex of the Moraceae family having a defensive
role against insects
Scientific
Plant name of the
family plant Category Compound Sub-category References
Ficus carica Protein Proteases Cysteine Ramos et al. (2010)
proteases
Moraceae Ficus elastica Serine proteases Patel et al. (2007)
and Tomar et al.
(2008)
Ficus carica Oxidase Peroxidase Saby et al. (2003)
(POD) and Sethi et al.
(2009)
Ficus carica Protease Serine Protease Azarkan et al.
inhibitors inhibitors (2004) and Kehr
(2006)
Antiaris Chemicals Cardenolide Toxicariosides Carter et al. (1997)
toxicaria
Morus Alkaloids Sugar-mimic Konno et al. (2006)
australis alkaloides

5.4 Different Associative Proteins

In addition to these PR proteins, many additional proteins and enzymes are also
present in the latex of Ficus carica leaves which may have a potential role in plant
defense mechanism. For example, lectins belonging to the carbohydrate-binding
protein group have been detected in the latex and are known to have anti-viral, anti-
bacterial, and anti-fungal roles (Macedo et al., 2015). Furthermore, though the study
on phosphatases is very few, some studies have shown the disturbance in the growth
and development of insect pests (Table 12.3) (Liu et al., 2011; Konno, 2011).

6 Role of Genotype and Environmental Factors in Ficus

Defense Mechanisms

Phenotypic variability consequences from interactions between genotype and envi-

ronment and is a first-rate driver of ecological and evolutionary interactions.
Measuring the relative contributions of a genetic variant, the environment, and their
interaction with phenotypic variation remains an essential intention of evolutionary
ecology. The genetic version is widely related to trait expression (Agrawal and
Hastings, 2019a; Geber & Griffen, 2003), supplying the variant on which choice
may also affect the duration of evolution. Environmental variation contributes to the
phenotypic version (Decker et al., 2019; Couture et al., 2015; Davis et al., 2005),
which influences specific genotypes to prosper (Beemelmanns & Roth, 2017; Potts
& Hunter, 2021).
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 299

6.1 Role of Genotype in Modifying the Defense Mechanism

The type of defense mechanisms in Ficus depends on several factors, such as an

individual organism, its variety, or species. However, the genotype of Ficus is the
most responsible for the differentiation of morphological and biochemical factors,
which include the color and concentration of latex; morphology of trichomes; size,
shape, location, and amount of chemical and mineral present.
The genetic elements are the significant player in determining the Ficus species
specific mineral system, which could elaborate the unique features of leaf minerals
(Pierantoni et al., 2018). As an example, the morphological features of phytoliths
could be an identifying factor for phylogenetic classification (Prychid et al., 2003).

6.2 Role of Plant Age and Season in Modifying

the Defense Mechanism

The organs in Ficus are not protected throughout the year equally as the aging and
occurrence of seasons have a major contribution towards altering the defense strat-
egy. It has been observed that the furanocoumarin content of F. carica leaves and
latex rises in the spring and summer but not in the autumn season (Zaynoun et al.,
1984). Marrelli and co evaluated the chemical composition, and concentration of
fruits of F. carica harvested in different months intervals, corresponding to distinct
phenological stages of the fig. The total amount of phenolics in figs increased as
they ripened, but the content of pyranocoumarin and furanocoumarin declined
(Marrelli et al., 2012). Another research discovered that the chemical content of dif-
ferent parts of F. carica (such as bark, leaves, and woody components) varies over
time and that these modifications are unique based on individual compounds and
organs (Marrelli et al., 2014). Moreover, the range or the amount of latex produced
varies according to the age of the plant part, i.e., young Ficus organs produce more
latex than older parts. Both seasonality and plant growth stages are expected to play
a major role in modulating the Ficus defense mechanism, and their effects are likely
to be coupled (Konno, 2011).

7 Inducible Defense Mechanism

Against insect herbivores, plants have developed a variety of defense systems.

These defense systems can be divided into two types: (1) constitutive or permanent
defenses and (2) inducible or transient (Karban & Baldwin, 1997). Both defenses
are achieved through comparable methods, but constitutive protections are inactive
form prior to an herbivore attack, whereas induced defenses or transient defenses
are activated only after an injury occurs (Gatehouse, 2002; Dicke et al., 2003; Chen,
300 S. Pattanayak et al.

2008). In addition to intrinsic defenses, initiating specialized herbivory-specific

defense responses is a key strategy for plant survival and persistence (Karban &
Baldwin, 1997).
Host plants can detect the type of injury or wound caused by mechanical means
or insects or herbivores to activate the type of induced defense strategy. Elicitors
like biotic (Insect, microbe), abiotic (Drought, UV, temperature, mechanical dam-
age, etc.), and chemical (Jasmonic acid, Salicylic acid, ethylene, metallic ion, salt,
etc.) are prerequisites for the induction of resistance response in plants (Fig. 12.3).
As a result, plants respond to many stimuli, including contact and the attacking
herbivore’s salivary enzymes. Physical and molecular mechanisms are involved in
this induction. Insect harm or mechanical damage to plants activates several signal
transduction pathways (Korth & Dixon, 1997). Sometimes, a minor alteration in the
host gene transcription can cause induced defenses. Although the genes involved in
this process differ between species, systemically triggered defenses arise due to
modification of the expression of genes (Walling, 2000). For example, photosyn-
thetic genes in Nicotiana tobacum are down-regulated in response to insect attack,
whereas defense genes are up-regulated (Hermsmeier et al., 2001). Induced resis-
tance mediated defense responses in plants are carried out in two broad aspects or
pathways, which include: – (i) direct defense against insects (secretion of inhibitory

Fig. 12.3 Induction of resistance responses concerning abiotic, biotic, and chemical elicitors
in Ficus
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 301

proteins, enzymes, and toxins) (ii) indirect defense (secretion of plant volatiles
attracting natural predator and parasites of harmful insects).
Physical and chemical factors are involved in the induction of defense systems.
Long-term consumption of plant parts by herbivores appears to strengthen the
toughness of Ficus leaf and results in a high rise in the density of calcium oxalate
crystal in a few species of Ficus (Xiang & Chen 2004). This herbivore injury can
lead to silicification or an increase in trichome density and number (Baur et al.,
1991; Massey et al., 2007; Strömberg et al., 2016; Liu et al., 2018) in some plants.
The expression of chemicals such as chitinase, trypsin inhibitor, and peroxidase can
be elevated in the latex of F. carica after several hours of wounding treatment (Kim
et al., 2003). However, several investigations on other plants have led us to believe
that herbivore or pathogen infection may affect the activation of the secretion of
specific Ficus defense chemicals. It was also reported that mechanical injury and
pathogen infection were shown to induce the synthesis of certain furanocoumarins
in parsnip (Krieger et al., 2018). Other factors may influence Ficus defenses, but
limited research on this area.

8 Evolutionary Trends Towards Complex Adaptation

The mechanisms through which complex adaptations develop are a fundamental

challenge in evolutionary biology. Complex adaptations are thought to take a long
time to develop because they require unique population genetic or environmental
variables to evolve. Complex adaptation blueprints in molecular systems, on the
other hand, are standard, implying that they can quickly evolve (Pal & Papp, 2017).
If complex adaptation of a plant is observed, then it would be found that each plant
organ has its unique histology and defensive architecture, and the amount of defen-
sive chemicals varies from one organ to the next (Villard et al., 2019). Insects and
pests coevolved with Ficus defense apparatus do so primarily through various tac-
tics to attack the host and intricate adaptability. Ficus are thus plants that shed light
on the exciting arms race those plants engage in to adapt to their surroundings.
Evolutionary trends toward complex adaptability are ongoing and will continue to
evolve in tandem with nature.

9 Evolution and Coevolution

Biotic stresses are one of the most powerful selective pressures on the plant, and
their interaction with the host assists in improving both organisms’ genetic diver-
sity. Host and pathogen selection toward each other is highly significant as if a
pathogen chooses its host and can adapt to the host system. Then the host also
imposes the same levels of behaviour or selection pressure on its host, resulting in
302 S. Pattanayak et al.

continuous mutual adaptation between them. Co-evolutionary interactions are

highly related to organism evolution.
Various generalist and specialist insects prey on Ficus species (Novotny et al.,
2010; Volf et al., 2018). It has been claimed that the generalist insects’ defense
mechanisms are often harmful compounds, resulting in higher mortality. Plants use
a variety of tactics to defend themselves, including chemical diversity pyramiding
and mechanical defenses, which reduce insect feeding efficiency and increase feed-
ing duration resulting in a long time of contact with toxic compounds and harmful
biotic agents, which lead to a high increase in insect mortality (Koricheva et al.,
2004; Volf et al., 2015, 2018). In 2018, the link between Ficus pests and several
Ficus defense features was explored, which showed the critical role of chemicals
like protease, alkaloid, polyphenol oxidative activities, and structural features such
as trichome length, trichome density.
On the other hand, in this insect-plant war, certain insects are smart enough to
overcome the host’s defense activities by secreting specific enzymes that can neu-
tralize these host chemical weapons (Volf et al., 2018). To avoid latex toxicity, cer-
tain insects discovered the piercing technique in canal branches resulting emission
of plant latex, freeing the distal tissues from toxic chemicals (Konno, 2011), after
which it is safer to consume the distal tissues. Though this activity is not examined
in Ficus, other latex-producing plants have been extensively discussed (Dussourd,
2017). The host plant and its corresponding parasites reciprocally affect each oth-
er’s evolution during their interaction period and ultimately lead to the convolution
of both the organisms.

10 Pattern of Defense Mechanism in Various Parts of Ficus

A wide range of biotic agents chooses their infection site according to their prefer-
ence and suitability. The pests attacking the leaf may be or not be the same as the
pests attacking the other parts. Similarly, the pests attacking fruits or floral parts
may be or may not be the same as other pests infecting the same host. The histology,
chemical constituent, and concentration are not the same for different parts of the
plant, so the infecting pests and their infection strategy also vary. Moreover, there is
no doubt that the mechanism of defense by the host in various parts also differs. For
example, the herbivores mainly target the foliar part of the plant, among other parts.
Similarly, ficin is the main PR protein in trunk and fruits, while trypsin inhibitors
are mostly present on the latex of leaves. Additionally, physical barriers such as
trichomes and a wide variety of chemicals and minerals help the host plant Ficus get
rid of herbivores. For example, Lignification, or the presence of lignin, is another
important physical barrier in figs plant that increases mechanical protection and
keeps the herbivores away from its trunk, bark, and other woody parts. The other
chemical factors that help in defense mechanisms are ficin, PR proteins, chitinase
acid phosphatases, furanocoumarins, wide antifungal, antibacterial, antiviral, insec-
ticidal, and insecticidal nematicidal activities (Kitajima et al., 2018).
12 Defense Mechanism of Fig (Ficus carica) Against Biotic Stresses… 303

A different set of defense mechanisms has been observed in different plant parts
dependent on the function of those plant parts. For example, the role of the insect
aganoid wasp and vertebrate species are essential for the pollination of figs and dis-
semination of seeds, respectively. Therefore, the toxic chemicals should be present
in an average concentration to not be more toxic to these host partners (Cruaud
et al., 2012; Kitajima et al., 2018).

11 Conclusion and Future Perspectives

For years tremendous research has been carried out to understand the defense mech-
anism strategy in various plants. The mechanism of defense highly varies from plant
to plant. The chemicals involved in the whole defense process are diverse and may
or may not be similar in all host plants. Similarly, the cascade of signaling to defense
mechanism varies among the members of the family Moraceae and different spe-
cies of Ficus. Moreover, the events of defense mechanisms are also highly varied
and different based on the plant’s diverse parts (leaves, stem, floral, etc.). These
strategies may be inducible or constitutive. Due to the new strain development in the
pest population, environmental function, and synergistic action of all mechanisms,
the host defense strategy has also evolved, resulting in broad resistance to various
pests and pathogens. One of the most studied defense strategies is the latex of the fig
plant, which can act as both mechanical and chemical means of defense strategy.
However, the concentration of latex and the size of the aggressor are highly related
as a small concentration of latex can be harmful to a tiny insect but may not be
harmful against a large size animal herbivore. Therefore, the Ficus group of plants
can provide some novel in-depth conclusions about the fascinating concept or strat-
egies for host adaptation to the external environment. Exploiting omics data like
genomes or transcriptomes of Ficus religiosa and Ficus carica which can be
accessed from public databases such as Genbank, can open up the path for new
advancements shortly.

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Part II
Fig (Ficus carica): Chemistry,
Functionality and Health-Promoting
Properties
Chapter 13
Chemistry and Nutritional Value of Fresh
and Dried Fig (Ficus carica)

Mohamed Fawzy Ramadan

1 Introduction

The fruit of the Ficus carica tree, which is part of the plant family Moraceae, is
sweet and nutritious (Solomon et al., 2006; Arvaniti et al., 2019; Khadivi &
Mirheidari, 2022). The Ficus carica genus contains more than 800 different kinds,
most of which are grown in regions with hot, dry climates like the Middle East and
the Mediterranean (Ouchemoukh et al., 2012; Harzallah et al., 2016; Meziant et al.,
2015; Solomon et al., 2006). Figs are a seasonal fruit that could be picked twice a
year, depending on the variety, either in the spring and summer or at the start and
end of the summer (Ouchemoukh et al., 2012; Vallejo et al., 2012; Arvaniti et al.,
2019). It is edible fresh, dried, jam, juice, and even in other forms (Solomon et al.,
2006; Hoxha & Kongoli, 2016; Harzallah et al., 2016; Arvaniti et al., 2019).
Figs include beneficial natural antioxidant substances like carotenoids, organic
acids, vitamin E, and phenolic compounds. Due to their antioxidant traits, phenolic
compounds are the most well-liked; they are also significant contributors to color,
flavor, and scent. Therefore, phenolic acids and flavonoids are two primary subcat-
egories of phenolic substances (Chang et al., 2016; Gharras, 2009; Shahidi &
Ambigaipalan, 2015; Arvaniti et al., 2019).
Different species of dried and fresh figs have undergone qualitative and quantita-
tive examination of their phenolic content (Solomon et al., 2006; Bachir et al., 2013;
Kamiloglu & Capanoglu, 2013; Bachir & Louaileche, 2015; Ajmal et al., 2016;
Harzallah et al., 2016; Maghsoudlou et al., 2017; Arvaniti et al., 2019). In addition,
researchers have investigated how phenolic component concentrations are affected

M. F. Ramadan (*)
Department of Clinical Nutrition, Faculty of Applied Medical Sciences,
Umm Al-Qura University, Makkah, Saudi Arabia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 313
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_13
314 M. F. Ramadan

by fruit color, fruit variety, harvest season, and drying techniques (Caliskan & Polat,
2011; Harzallah et al., 2016; Konak et al., 2017; Pereira et al., 2017; Ersoy et al.,
2017; Slatnar et al., 2011; Arvaniti et al., 2019).
The primary goals of this work were to summarize the levels of phenolics and
carotenoids in fresh and dried figs and to present the main components and phyto-
chemicals in figs.

2 Nutrients and Phytochemicals in Fresh and Dried Figs

Figs, both fresh and dried, are a good source of amino acids, vitamins (thiamin and
riboflavin), and trace minerals (iron, calcium, and potassium) (Solomon et al., 2006;
Ouchemoukh et al., 2012; Viuda-Martos et al., 2015). In addition, figs are high in
fiber and antioxidant components but free of sodium, fat, and cholesterol (Solomon
et al., 2006; Viuda-Martos et al., 2015; Veberic et al., 2008; Arvaniti et al., 2019).
Figure 13.1 shows fresh and dried figs’ chemical makeup and nutritional value. It is

Fig. 13.1 Composition and nutrient content (value/100 g) of fresh and dried figs. (Source: Data
from the USDA National Nutrient Database for Standard References 2018 and Arvaniti et al., 2019)
13 Chemistry and Nutritional Value of Fresh and Dried Fig (Ficus carica) 315

evident that the chemical composition of fresh and dried figs differs, particularly in
terms of the amounts of total calories (kcal), sugars (g), calcium (mg), phosphorus
(mg), magnesium (mg), and potassium (mg).
Due to a large number of phytochemicals present in the fruit tissues, peel (skin),
and pulp, Ficus carica phytochemistry is complex. The fruits of the Ficus carica
tree contain a variety of phytochemicals, such as phenolics, anthocyanins, and
carotenoids (Jagtap & Bapat, 2019). Therefore, consuming whole, ripe fruits are
advised because fruit skins are frequently the primary source of phenolic chemicals
(Vinson et al., 2005).
Studies have reported the phytochemicals content of dried and fresh Ficus carica
entire fruits. Individual phenolic compounds and carotenoids, as well as total phe-
nolics (TPC), total flavonoids (TF), and total anthocyanins (TA) have been identi-
fied. The majority of the research showed that the phytochemical profile is
substantially influenced by the fig variety as well as by other elements like color,
fruit portion, level of maturity, and method of drying (Solomon et al., 2006;
Ouchemoukh et al., 2012; Yemis et al., 2012; Nakilcioglu & Hısıl, 2013; Bachir
et al., 2013; Bachir et al., 2014; Debib et al., 2014; Kamiloglu & Capanoglu, 2015;
Bachir & Louaileche, 2015; Bachir et al., 2016; Harzallah et al., 2016; Pereira et al.,
2017; Amessis-Ouchemoukh et al., 2017; Konak et al., 2017; Ersoy et al., 2017;
Sedaghat & Rahemi, 2018; Arvaniti et al., 2019).
TPC levels in fresh fig fruit ranged from 19 mg GAE/100 g fresh weight (FW) to
6147 mg GAE/100 g FW (Caliskan & Polat, 2011; Harzallah et al., 2016; Arvaniti
et al., 2019). TPC concentrations in dark-colored fig types ranged from 53 to
6147 mg GAE/100 g FW (Solomon et al., 2006; Caliskan & Polat, 2011; Harzallah
et al., 2016; Arvaniti et al., 2019). It has been demonstrated that fig peels contain
significantly more phytochemical components than fig pulp. In addition, it is com-
monly acknowledged that fruit and vegetable skin contains most anthocyanins, con-
tributing to their high phytochemical content and nutritional worth (Solomon et al.,
2006; Arvaniti et al., 2019).
Phenolic acids, flavonoids, and carotenoids are abundant in dried Ficus carica.
TPC of dried fig extracts has been estimated to be between 45 and 644 mg GAE/100 g
dry weight (DW) for light-coloured dried Ficus carica types and between 399 and
756 mg GAE/100 g DW for dark-colored dried Ficus carica varieties (Bachir et al.,
2013; Debib et al., 2014; Bachir & Louaileche, 2015; Arvaniti et al., 2019).
Additionally, studies have shown that anthocyanin pigments, such as peonidin,
cyanidin, petudinin, delphinidin, and malvidin, are primarily found in purple, red,
and other dark-coloured fruit (Khoo et al., 2017; Wang et al., 2017; Arvaniti
et al., 2019).
316 M. F. Ramadan

3 Specific Phytochemicals in Fresh and Dried Figs

Russo et al. (2014) assessed the phenolic composition of dried and fresh figs (culti-
var Dottato) from various local origins and contrasted the phenolic content of fresh
figs with dried figs from various origins (Turkey, Italy, and Greece). The Cilento
figs’ phenolic content was decreased by sun-drying. Therefore, to prevent an exces-
sive loss of phenolics, it is necessary to carefully control all the parameters involved
in fig drying, especially the amount of time the fruit takes is exposed to the sun and
other variables like environmental humidity.
The most common substances in dried and fresh Ficus carica are phenolic acids,
including gallic acid, ferulic acid, syringic acid, chlorogenic acid, caffeic acid, and
cinnamic acid. Gallic acid (5 mg/100 g DM) and chlorogenic acid (32 mg/100 g
DM) are the two phenolic acids that are most prevalent in dried Ficus carica,
although the same compounds were detected in fresh Ficus carica with quantities of
6.9 mg (gallic acid)/100 g DM and 28 mg (chlorogenic acid)/100 g DM, respec-
tively (Arvaniti et al., 2019).
The highest concentrations of rutin, quercetin-3-O-rutinoside, and epicatechin
were 14.6, 10.8, and 36.6 mg/100 g DM in dried Ficus carica. These substances
accounted for 68 mg/100 g DM for rutin, 38 mg/100 g DM for quercetin-3-O-
rutinoside, and 76.9 mg/100 g DM for epicatechin in fresh fig (Arvaniti et al., 2019).
The two primary anthocyanin pigments in all Ficus carica cultivars were
cyanidin-3-O-rutinoside and cyanidin-3-O-glucoside. In dried and fresh Ficus car-
ica, cyanidin-3-O-rutinoside concentrations varied between 73.0 and 91.0 mg/100 g
DM (Arvaniti et al., 2019). Proanthocyanidins, in particular, are found in high con-
centrations in figs (Vinson, 1999; Russo et al., 2014), wherein the cultivar and time
of harvest are just two examples of the many variables that affect the concentration
of these compounds (Burda et al., 1990; Kennedy et al., 2001; Crisosto et al., 2010;
Russo et al., 2014).
Both dried and fresh figs have been found to contain lutein, beta-carotene, zea-
xanthin, and beta-cryptoxanthin. However, fresh figs were shown to have the most
concentrations of beta-carotene, zeaxanthin, and lutein, whereas dried figs had the
most concentrations of beta-cryptoxanthin and lutein (Arvaniti et al., 2019).

4 Future Perspectives

This work reported the nutrients, and phytochemical composition of Ficus carica
dried and fresh figs. Figs, both fresh and dried, are a great source of bioactive phy-
tochemicals. Topics of particular concern are how figs’ harvesting practices and
packing techniques affect the fruits’ phytochemical makeup and antioxidant capa-
bility. In addition, it is crucial to look into alternative drying methods (such as
freeze-drying and microwave-drying) for dried figs to preserve their phytochemical
content and lengthen their shelf life.
13 Chemistry and Nutritional Value of Fresh and Dried Fig (Ficus carica) 317

To recommend the ideal circumstances and processes for Ficus carica, it is

essential to investigate further the impact of various drying techniques on the phy-
tochemical levels and nutritional value of Ficus carica and environmental factors.
Furthermore, the potential uses of Ficus carica bioactive compounds and extracts as
functional constituents of food items should be investigated in light of the existence
of phytochemical substances in Ficus carica phytochemicals and their association
with the bioactive potential. Finally, future clinical and dietary studies might help us
better comprehend figs’ positive impacts on human health.

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Chapter 14
Fig Seeds: Source of Value-Added Oil
Within the Scope of Circular Economy

Lahcen Hssaini

Abbreviations

ABTS 3-ethylbenzothiazoline-6-sulfonic acid

ALA alpha-linolenic acids
CE circular economy
DPPH 2,2-diphenyl1-picrylhydrazyl
DR desaturation ratio
EFAs essential fatty acids
FA(s) fatty acid (s)
FSO fig seeds oil
GAE gallic acid equivalent
IC50 half maximum inhibitory concentration
IV iodine value
LA linoleic acid
LCPUFA long chain polyunsaturated fatty acids
LDR linoleic desaturation ratio
MUFA monounsaturated FA
ODR oleic desaturation ratio
PUFA polyunsaturated FA
SCPUFA short chain polyunsaturated fatty acids
SV saponification value
TSFA total saturated FA
TUFA total unsaturated FA

L. Hssaini (*)
National Institute of Agricultural Research, Avenue Ennasr, BP 415 Rabat Principale,
Rabat, Morocco
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 321
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_14
322 L. Hssaini

1 Fig Seeds: A Novel Source of Oil

Figs (Ficus carica L.) are whispered to be the oldest of all cultivated fruit species
and the first domesticated tree of the Neolithic Revolution (Kislev et al., 2006).
Since then, figs have been a major food source and have been utilized alongside
other tree parts such as leaves, latex, bark, and roots for medicinal purposes
(Badgujar et al., 2014). Due to its multiple health-promoting properties and biologi-
cal activities, figs invite the attention of the scientists worldwide and have been
widely studied, as a whole, for their biological compounds and activity along with
their antidiabetic properties (Debib et al., 2014; Del Caro & Piga, 2008; Hssaini
et al., 2019; Jeong & Lachance, 2001; Kamiloglu & Capanoglu, 2013; Schmitzer
et al., 2011; Veberic et al., 2010). However, seeds remain the less studied part of the
species (Hssaini et al., 2020b, 2021b). Thus, very few reports have been undertaken
on their lipochemical and nutraceutical properties (Ben-Othman et al., 2020). Figs
produce a considerable amount of yellowish and round-shaped seeds, with numbers
fluctuating from 30 to 1600 and an average thousand seed weight of about 1.14 g
equivalent (Hssaini et al., 2020a). The formation of seeds is dependent on the mutu-
alism between the tree and the wasp pollinator. Therefore, the seeds are generally
hollow without receiving the caprifig (male tree) pollen on the female of blastoph-
aga psenes body (Kato & Kawakita, 2017). These seeds are known to contribute to
the fruit’s taste and flavor along with its health-promoting attributes (Hssaini et al.,
2020a; Nakilcioğlu-Taş & Ötleş, 2021). Only very few studies have been interested
in exploring fig seeds’ lipochemical and bioactive compounds (Badgujar et al.,
2014; Hssaini et al., 2020a, 2021b; Jeong & Lachance, 2001). In the studies of
Jeong and Lachance (2001), Hssaini et al. (2020a, 2021a), results showed that fig
seeds account for approximately 20 to 30% of oil (Hssaini et al. 2020a, 2021b),
which is were comparable to oil content levels found in other fruits seeds such as
kiwi (28.3%) (Taylor et al., 2011), papaya (27.87%) (Senrayan & Venkatachalam,
2019), pumpkin (27.83-%) (Wang et al., 2018), Mangosteen (21.68%) (Ajayi et al.,
2007) and honeydew (25%) (Yanty et al. 2008). These yield levels are much higher
than those found in prickly pear seeds, which exhibited an oil content of 5.4–9.9%
(Chougui et al., 2013; Mannoubi et al. 2020), guava (12.6–16%) (Piombo et al.,
2006; Prasad & Azeemoddin, 1994), grape (8.66–13.6%) (Lucarini et al., 2020;
Tangolar et al., 2009) (Fig. 14.1).
As the first macro quality component of interest for all fruit seeds, the oil yield-
ing is mainly dependent on the varietal factor, geographical variations, or pollen
source (Raihana et al., 2015). In the first report of Hssaini et al. (2020a) on seeds
extracted from four fig cultivars in Morocco, the impact of the phenotypic factor
was highly significant on oil yield and chemical attributes. Authors attributed the
large variation in those variables to the cultivar factor, as the sampled figs belonged
to an ex-situ collection conducted under the same condition. In the same way, sev-
eral studies have indicated that the genetic factor has a greater influence on seed oil
yield and quality (Górnaś & Rudzińska, 2016). With a highly promising oil yield,
fig seeds have all the potential to be economically attractive for food and non-food
14 Fig Seeds: Source of Value-Added Oil Within the Scope of Circular Economy 323

Fig. 14.1 Oil yield (%) of several fruit seeds compared to figs. Orange color indicates high oil
yield, while turquoise refers to yield level. The low color intensity means the lower value and
vice versa

applications, similar to other fruits’ seeds oil such as grape, pomegranate, guava,
honeydew, and kiwi (Górnaś & Rudzińska, 2016; Hssaini et al., 2020a; Piombo
et al., 2006; Raihana et al., 2015). Besides, encouraging bringing this novel oil to
the worldwide markets at favorable prices could lead to a tremendous economic
impact for communities where this species is abundantly growing and significantly
contribute to the household economy.

2 Fatty Acids Profile

As a minor vegetable oil from atypical sources, fig seeds oil (FSO) was less docu-
mented, and therefore very few data are available on this concern. Fig seeds produce
a light yellowish colored oil with a high unsaturation rate reaching up to 88%, with
a remarkable phenotypic effect (Hssaini et al., 2020a; Hssaini et al., 2021b).
Linolenic (C18:3) and linoleic (C18:2) acids are the major fatty acids in FSO pres-
ent in the following ranges 38.4–43.5% and 28.9–34.5%, respectively (Table 14.1).
Oleic acid (C18:1) is the third predominant fatty acid (FA), with rates varying
between 13.4 and 15.6% (Hssaini et al., 2020a). High levels of unsaturated fats are
324 L. Hssaini

Table 14.1 Fatty acid profiles of fig seed oil in comparsion with other fruit seed oils
Source of Prickly
seeds Fig grape honeydew Kiwi Guava pear Durian
Oil content 21–30 – 13–37 26.8– 12.6–16 5.4–9.9 0.5–0.5
(%) 28.3
Fatty acids – – – – – – –
(%):
C12:0 – 0.24– 0.14–0.23 – – – –
0.24
C13:0 – – 0.06–0.24 – – – –
C14:0 – 0.03–0.3 0.07–0.25 – 0.1–0.2 0.11–0.11 –
C15:0 0.02–0.02 – 0.03–0.13 – – – –
C15:1 – – 0.08–0.31 – 6.6–7.8 – –
C16:0 8.5–9.11 8–11.55 8.51– 5.72– – 12.76–26.8 3.5–9.7
23.88 8.02
C16:1 0.06–0.17 0.07– 0.08–0.1 – – 0.75–2.01 0.21–
0.39 0.3
C17:0 0.03–0.07 0.03– 0.08–0.19 – – – –
0.14
C17:1 0.04–0.07 – 0.02–0.03 – – – –
C18:0 2.51–3.3 4.24– 4.89–6.09 3.2– 4.5–5.8 3.2–4.32 3–14
4.96 4.55
C18:1 13.32– 18.14– 0.1–31.5 14– 8.2–10.8 16.41–19.3 7–26.2
15.66 23.09 18.79
C18:2 28.85– 59.02– 51.6–61.1 14.08– 76.4–77 49.7–60.69 44–
34.56 67.34 18.43 80.07
C18:3 38.43– 0.21– 0.19–0.87 51.2– 0.1–0.3 1.56–3.9 0.1–
43.61 1.04 61.41 0.13
C20:0 0.02–0.17 0.05–0.4 0.22–0.29 – 0.3–0.5 0.3–0.3 –
C20:1 0.04–0.2 0.03– 0.08–0.1 – – – –
0.15
C20:2 – – – – – – –
C22:1 – – – – – – –
C24:0 – – – – – – –
DR 4.93–5.04 2.6–3.69 1.96– 4.25– 7.16–9.33 2.78–3.79 3.06–
297.64 4.66 6.3
LDR 0.87–0.91 0.01 0.01 1.65– – 0.02–0.06 –
1.82
ODR 0.83–0.83 0.72– 0.66–1 0.81– 0.88–0.9 0.74–0.790.75–
0.79 0.82 0.86
ω-6/ω-3 0.75–0.79 56.75– 70.15– 0.28– 256.67– 12.74–38.9 440–
320.67 271.58 0.3 764 615.92
MUFA 13.45–16.1 18.36– 0.28– 14– 8.2–10.8 17.16– 7.21–
23.51 31.73 18.79 21.31 26.5
PUFA 67.27– 60.06– 51.79– 65.28– 76.5–77.3 53.6–62.25 44.1–
78.17 67.55 61.97 79.84 80.2
(continued)
14 Fig Seeds: Source of Value-Added Oil Within the Scope of Circular Economy 325

Table 14.1 (continued)

Source of Prickly
seeds Fig grape honeydew Kiwi Guava pear Durian
TUFA 80.72– 83.57– 52.07– 79.28– 84.7–88.1 74.91– 51.31–
94.27 85.91 93.7 98.63 79.41 106.7
TSFA 11.07– 13.07– 13.95– 8.92– 4.9–6.5 16.37– 12.7–
12.67 16.87 31.06 12.57 31.12 17.5
MUFA/PUFA 0.18–0.22 0.27– 0.01–0.51 0.21– 0.11–0.14 0.28–0.4 0.16–
0.39 0.24 0.33
TSFA/TUFA 0.13–0.14 4.95– 0.27–0.33 0.11– 0.06–0.07 0.21–0.42 0.16–
6.57 0.13 0.25
Iodine value 76.26– – 97.1–97.1 – 13.4–13.4 107.38– –
(IV) (g/100 g 76.63 107.38
oil)
Saponification 201.49– – 116–116 – 196–196 173–173 –
value (SV) 201.67
(mg KOH/g)
References Hssaini Tangolar Yanty Piombo Narváez- Ennouri Ho and
et al., et al. et al. et al. Cuenca et al. Bhat
(2020a, (2009); (2008); (2006); et al. (2005); (2015);
2021b) Lucarini Mallek- Wang (2020); Karabagias Mohd
and Jeong et al. Ayadi et al. Kapoor et al. Ali
and (2020) et al. (2018) et al. (2020) et al.
Lachance, (2018) (2020) (2020)
(2001)
MUFA Monounsaturated fatty acids, PUFA Polyunsaturated fatty acids, TSFA Total saturated fatty
acids, TUFA Total unsaturated fatty acids

%C 20 : 1 %C 22 : 1
DR desaturation ratio ;
%C 20 : 1 %C 22 : 1 %C18 : 1 %C18 : 2 %C18 : 3
%C18 : 1 %C18 : 2
ODR oleic desaturation ratio
%C18 : 1 %C18 : 2 %C18 : 3

%C18 : 3
LDR linoleic desaturation ratio ; 6 / 3 18 : 2 / 18 : 3
%C18 : 2 %C18 : 3

important for the balance in the human immune system as it has a natural defensive
function against cardiovascular diseases and in mitigation of some other health
issues (Chougui et al., 2013; Ramadan 2019). Thus, C18:3 plays a significant role
in reducing inflammatory disorder’s severity, whereas C18:2 is involved in enhanc-
ing some physiological functions, including immune response and blood clotting,
due to its ability to convert to eicosanoids hormone. Furthermore, C18:1 has a role
in lowering plasma cholesterol (Gonçalves-de-Albuquerque et al., 2016; Park et al.,
2018). Unlike other fatty acids, Linoleic acid (C18:2n-6) and α-linolenic acid
(C18:3n-3) are essential FAs (EFAs) that must be regularly consumed through food,
including oilseeds and oil-bearing crops, since they are not synthesized in the body
326 L. Hssaini

and a deficiency may leads to transepidermal water loss increases, skin dryness,
nails crack as well as hair-loss (Sassa & Kihara, 2014; Vermaak et al., 2011).
Regarding saturated fats, palmitic acid (C16:0) was predominant in FSO within a
narrow range of variation of 8.54 and 9.05%. Stearic acid (C18:0) was the second
major saturated FA ranging from 2.50 to 3.30% (Table 14.1). According to Hssaini
et al. (2020a), pentadecylic (C15:0), palmitic (C16:0), palmitoleic (C16:1), marga-
ric acid (C17:0), arachidic acid (C20:0) and gondoic acid (C20:1) were only identi-
fied in minor levels, generally between 0.8 and 0.2% (Table 14.1). The same authors
reported highly significant differences among sampled fig trees based on all FAs,
with the exception of margaric, arachidic and pentadecylic acids besides the
TSFA. Looking at the lipid profile, FSO seemed fairly similar to the oil extracted
from kiwi seeds, which displays an unsaturation rate in the range of 79.2 and 98.6%,
which is comparable to FSO unsaturation rate that falls in the range of 80.7 and
94.2%. Besides, C18:3 and C18:2 are the major FAs in both species seeds followed
by oleic acid (Table 14.1). Unlike FSO, oil extracted from seeds of grape, honey-
dew, guava, prickly pear and durian have high levels of monounsaturated fatty acids
(MUFA), a fraction dominated by C18:1 (59.0–67.3%, 51.6–61.1%, 76.4–77%,
49.7–60.6% and 44–80.0%, respectively) (Table 14.1), which is rather similar to
olive oil. It is noteworthy that no data were reported on the following FA C20:2,
C22:1, C24:0.

3 Fatty Acids Ratios

Assessment of seeds from different phenotypes and highlighting any potential

diversity is not eventual for minor oil-bearing crops breeding. According to Velasco
et al. (1998), emphasizing the lipchemical diversity through fatty acid profiling of
considerable sample length of oil-bearing crops is difficult because they are inter-
correlated, and any intensive selection or breeding modification may significantly
affect the whole system. For this sake, a set of several FAs ratios, such as elongation
ratio (ER), desaturation ratio (DR), oleic desaturation ratio (ODR), and linoleic
desaturation ratio (LDR), are usually used to overcome this issue. The ER ratio
evaluates the relative weight of the elongation pathway in the chain length of mono-
unsaturated FA, from oleic acid (C18:1) to eicosenoic and erucic acids (C20:1and
C22:1, respectively). Desaturation ratio (DR) describes the efficiency of the desatu-
ration pathway in the chain length of polyunsaturated FAs. Thus it estimates the
weight length in elongation from C18:1 to C18:2 (ODR) and from C18:1 to C18:3
(LDR) within the overall FAs biosynthetic system. It is noteworthy that the FAs
desaturation cycle is mainly driven by genetic and environmental factors, including
temperature (Clemente & Cahoon, 2009; Sakamoto & Murata, 2002). The overall
n-3 and n-6 polyunsaturated FAs (PUFA) pathways are summarized in Fig. 14.2,
where several enzymes are involved in chain elongation and desaturation. The
desaturation efficiency in seed oils is essential since it has an important role in sev-
eral metabolism functions, including immune system stability (Chougui et al., 2013;
14 Fig Seeds: Source of Value-Added Oil Within the Scope of Circular Economy 327

Fig. 14.2 n-3 and n-6 fatty acids pathway desaturation and elongation. The enzymes ∆6 and ∆5
desaturases are encoded by FADS2 and FADS1, respectively (Brayner et al. 2018)

Ramadan, 2019). Growing evidence suggests that increasing dietary intake of n-6
and n-3 long-chain polyunsaturated fatty acids (LCPUFA) has significant effects on
neuronal metabolism by lowering neuroinflammatory processes and enhancing syn-
aptic plasticity and neurogenesis. In FSO, higher desaturation efficiency in the
LCPUFA pathway was reported, explaining the significant increase in the desatura-
tion rate recorded in all investigated samples. Thus DR, LDR, and ODR values were
in the range of 4.93–5.04, 0.87–0.91, and 0.83–0.83, respectively, which is quite
similar to the kiwi seeds oil desaturation efficiency (4.25–4.66 and 0.81–0.82 for
DR and ODR, respectively), with the exception of LDR ratio that lower in FSO
compared to kiwi seeds oil (1.65–1.82). It is noteworthy that, based on the data sum-
marized in Table 14.1, FOS seemed to exhibit high efficiency in LCPUFAs path-
ways compared to the fruits’ seeds listed in Table 14.1. However, guava seeds oil
appeared to display high levels for both DR (7.16–9.33) and LDR (0.8–0.9) ratios
compared to FSO. Besides FAs desaturation ratios, the balance between omega-6
(ω-6) and omega-3 (ω-3), represented by linoleic acid (LA) and α-linolenic acids
(ALA), respectively, constitutes an important determinant of seeds oils quality and
is a critical component in human diet (Simopoulos & Gene, 2016). Generally, FSO
displays a very low ω-6/ω-3 ratio which falls in the range of 0.75 and 0.79. Due to
competing functions of both n-6 (LA) and n-3 (ALA) PUFAs in the formation of
highly proinflammatory eicosanoids, balanced intake of LA and ALA FAs is
required to avoid chronic diseases and to uphold good health (Saini & Keum, 2018).
It is worth mentioning that nutrition societies acclaim a lower value of ω-6/ω-3 ratio
for a balanced and healthy diet. Compelling evidence advises that low levels in the
balance between ω-6 and ω-3 is significantly associated with reducing the risk of
chronic cardiovascular disease and enhancing neuroinflammation alongside
328 L. Hssaini

protecting and repairing brain cells protection (Dyall. 2017; Saini & Keum, 2018).
Therefore, the intake of fig seeds oil may be very beneficial for human diet as it
presents a very low ω-6/ω-3 ratio (0.75–0.79) (Table 14.1) compared to several
vegetable oil sources, since the ratio in most of western societies is overall as high
as 15:1. For blackberry seeds this ratio (3.63) was reported to be three times higher
than that in elderberry seed oil (1.19) (Domínguez et al., 2020). In other berries such
as bilberry, cranberry, gooseberry, and strawberry this ratio was reported to be close
to 1.0 (Alves et al. 2020). In the fruits’ seeds summarized in Table 14.1, all species
exhibited very high levels of ω-6/ω-3 ratio, with the exception of the kiwi seeds oil,
which seems once again very close to the FSO in terms of FAs profile.

4 Chemical and Biochemical Properties

Unfortunately, fig seed oil biochemical properties have been less documented and
almost gone unheeded. As far as we know, the only available report on this aspect is
the one carried out by Hssaini et al. (2020b), where the results are summarized in
Table 14.1 and Fig. 14.3. Iodine value (IV) showed a significantly narrowed varia-
tion range of 76.2–76.6 g/100 g of fig seeds oil, classifying it as non-drying oil,
owing to the low IV determined in fig seeds from four cultivars. These IV levels
indicate the high unsaturation degree found in FSO fats. It is noteworthy that IV
determines the number of mg of iodine required to saturate the fatty acids through
the ability of an unsaturated carbon to carbon bond (Anang et al., 2019).
Saponification value (SV) displayed high levels of 201, suggesting a high amount of
FA with low molecular weight. Thus, glycerides made of short-chain polyunsatu-
rated FAs (SCPUFA) exhibit higher SV than LCPUFA. SV indicates oil alkali-
reactive groups (Cerchiara et al., 2010), which helps predict the type of glycerides
in fats. According to Table 14.1, IV of honeydew, guava, and prickly pear oils are
97.1–97.1, 13.4–13.4, and 107.3–107.3 g/100 g of oil, respectively. The same seed
types exhibited PV levels in 116–116, 196–196, and 173–173 mg KOH/g. Among
the latter, guava seemed to show comparable physicochemical attributes to
FSO. Furthermore, compared to FSO, rambutan seed oil showed IV in 41.8–49.6 g
I2/100 g and an SV ranging from 157 to190 mg KOH/g, respectively (Leong &
Shui, 2002).
Primary and secondary oxidation of FSO evaluated through ultraviolet (UV)
absorbency at 232 and 270 nm suggests the occurrence of some hydrolytic reactions
during the oil chemical extraction (method 659:199) (Hssaini et al., 2021b). UV
absorbance at the following wavelengths, 232 and 270 nm, were in the range of
1.83–5.36 and 0.21–0.7, respectively, showing a significant impact of the cultivar
factor (Fig. 14.3). The only available data regarding FSO total phenolics, antioxidant
activity, and IC50 are reported by Hssaini et al. (2020a), summarized in Fig. 14.3.
Total phenolics were reported in 69.8 and 100 mg GAE/100 g of oil. These amounts
are comparable with those found in prickly pear (48–89 mg GAE/100 g) (Chougui
et al., 2013). However, they are higher than levels reported in other fruit seed oils
14 Fig Seeds: Source of Value-Added Oil Within the Scope of Circular Economy 329

Fig. 14.3 Violin and box plots demonstrate the density of the scores within variables related to the
biochemical properties of FSO. Total phenols (mg GAE/100 g), DPPH· and ABTS antioxidant
activity (mg Trolox equivalent/g oil) and the half-maximum inhibitory concentration IC50 (mg/
mL). The embedded graph at the top right shows the ultraviolet oil absorbance at 232 and 270 nm

like mango, avocado, and jackfruit (1.17, 0.88, and 27.7 mg GAE/100 g, respec-
tively) (Wang et al., 2018). Furthermore, FSO exhibited high antioxidant potency
through DPPH· (2,2-diphenyl1-picrylhydrazyl) and ABTS°+ (3-ethylbenzothiazo-
line-6-sulfonic acid) free scavenging assays. Available results showed interesting
antioxidant activity ranging from 226.4 to 294.3 and 136.6 to 22.65 mg Trolox
equivalent/g oil for DPPH· and ABTS assays, respectively (Fig. 14.3). Both assays’
half-maximum inhibitory concentration (IC50) was 19.2–68.7 and 17.4–44.8 mg/
mL, respectively. It is noteworthy that the aforementioned results depend on the
phenolic profile more than phenols amounts, owing particularly to the fact that sam-
ples extracts contain a mixture of different phenolic compounds, making it difficult
to attribute the radicals scavenging ability to one or a few active principles (Wu
et al., 2009). Owing to data reported above, FSO is found to be rich in phenolics and
present high free radical scavenging activity even more important than what have
been reported over prickly pear seeds oil (Chougui et al., 2013).
All biochemical attributes aforementioned strongly suggest that FSO can be a
potential natural source of antioxidants for the food industry and pharmaceutical and
cosmetic applications. However, no single oil can accomplish all the requirements
and standards of the growing oil sector. Since there has been a renewable nutritional
interest for unsaturated oils with higher antioxidant capacity. Consequently, Further
investigations on other aspects such as FSO vitamins, phenolics, and volatile com-
pounds are required to evaluate their potential applications as an atypical source of
highly nutritive oil, functional food ingredients, additives, or feed.
330 L. Hssaini

5 Fig Seeds Valorization Through Circular Economy

Today, the global food system is under severe stress owing to inequities in produc-
tion and chain value about the misuse and over exploitation of natural and man-
made resources, which are highly vulnerable to ongoing climate change. Nowadays,
it is a common belief that there is an urgent need to increase the sustainability and
resilience of our food systems to meet the demand of the exponentially growing
population. Therefore, the global food production major challenge is to increase by
at least 60% by 2050, as the world population is expected to reach 9.7 billion people
(European Environment Agency (EEA), 2016). In this context, the circular econ-
omy (CE) has gained traction in the scientific community as a pioneer for innova-
tive, resilient, and sustainable food systems (Ancut, 2020). CE embodies an ultimate
alternative to the conventional linear take-make-consume-dispose economic model.
According to Steffen et al. (2015) and Fidelis et al. (2019), the CE is an umbrella
concept aiming at reducing waste generation and minimizing the use of new inputs
and energy while mitigating the pressures on the environment.
Fruits processing, like other agriproducts valorization systems, generates consid-
erable amounts of waste, which is a challenging endeavor that, if not accurately
addressed, may negatively impact the environment and the economy alongside the
ability to meet the population’s needs in new food sources. Daily, the entire food
system loses about 30% of all produced foods, where the mass of by-products gen-
erated by the fruit processing industry is becoming higher among all types of agri-
foods, reaching up to more than 190 million tons of by-products worldwide (Campos
et al., 2020; Gençdağ et al., 2020). In the fruit sector, more than 45% of the total
production is wasted during the post-harvest valorization chain; For western Asia
and northern Africa, this loss is about 11% (FAOSTAT, 2021). Seeds constitute a
relevant part of this waste, as being usually discarded after fruit processing to jam,
juice, syrup, or must (Ben-Othman et al., 2020). Therefore, there is an urgent need
to transition from current linear food production to a circular economy model to
recycle these seeds and recover resources for high-value-added products (Ancut,
2020; Campos et al., 2020; Fidelis et al., 2019; Lucarini et al., 2018). Large seeds
generated from the conventional linear processing model are not yet appropriately
exploited for figs. Thus, various potential products from this waste, particularly oil,
press cake, or meal, are not yet systematically available on the market or perhaps
not easily marketed as common oilseeds used by the food industry (i.e., sesame,
sunflower, and soybean), that have been promoted for a long time (Alves et al.,
2020; Ramadan, 2019).
Unfortunately, there is a severe lack of consistent statistics regarding waste
amounts generated during fig processing, including peels, latex, and seeds. On the
contrary, for other fruits, like mangos and oranges, available reports stressed a waste
level in a range of 30–50%, while it can be in a higher range in the case of durian,
jackfruit, and mangosteen, which can generate up to 70% of waste (Fidelis et al.,
2019). Figure 14.4 describes the linear model of fig processing, explores potential
products and byproducts produced, and proposes a circular model by reprocessing
14 Fig Seeds: Source of Value-Added Oil Within the Scope of Circular Economy 331

and reusing wastes for some potential industrial applications. Generally, figs are
harvested for fresh consumption or transformation at the full ripening stage or left
on the tree until they shrivel or become half-dried. In some cases, figs can also be
harvested at their full maturity stage and then dried with industrial ovens or dryers
with appropriate pretreatments to prevent browning. Dried figs, as a whole or sliced,
can be roasted and used as a coffee substitute (Ishurd et al., 2004; Liang et al., 2016;
Mat Desa et al., 2019) or used to produce wine and spirits, with either plurivarietal
or monovarietal dried figs as the raw inputs (Solana et al., 2018).
Being a highly perishable commodity, a huge part of harvested fresh figs are
processed into jam, syrup, juice, wine, or must. During which seeds received no
intention and almost underwent unheeded. Seeds are usually crushed during pro-
cessing, and thus their potential importance goes unnoticed or is often discarded as
solid waste. As previously stressed, these seeds can be exploited as an essential
source of highly unsaturated oil with high antioxidant potency. Owing to FSO lipo-
chemical and biologic properties, it can be brought to the international markets at
favorable prices (Hssaini et al., 2021b) as it can be successfully used for food and
cosmetic applications. During oil extraction, press-cakes or meals are produced
from which can be extracted high-value components such as fats, antioxidants, pro-
teins, dietary fiber, and phytochemicals. They can also be used as fiber-based foods
and dietary supplements to formulate new products or for animal feed (Fig. 14.4).
So far, available literature has proposed several uses of fruit seeds byproducts in
industrial formulations, such as muffins and cookies enriched with fruits seeds such
as grape seeds and date seeds (Karnopp et al., 2015, 2017; Maqsood et al., 2020).
These studies, among others, have shown that the incorporation of fruit seeds in
newly formulated products resulted in enhancing their nutritional virtues (Bouaziz

Fig. 14.4 Global diagram of fig processing and circular economy of waste materials focused
on seeds
332 L. Hssaini

et al., 2010; Platat et al., 2015). In the proposed CE, we could also spot other wastes,
solid like peels and liquid such as latex, which can be an important source of essen-
tial bioactives (pigments, proteins, enzymes, dietary fibers, etc.) for many industrial
applications. This waste is reported in several previous studies as potential nutra-
ceutical and functional food ingredients (Backes et al., 2018; Barolo et al., 2014a,
b; Hssaini, et al., 2021a; Milošević et al., 2020; Sirisha et al., 2010). Figure 14.4
provides a holistic overview of the fig processing pathways, focusing on seeds valo-
rization as they remain the less studied and exploited part of the species. It is note-
worthy that all valorization pathways of fig products and byproducts fundamentally
depend on the egentic factor.

6 Conclusions and Future Research Directions

Given the continuous progress of the oilseed industry, an important interest is cur-
rently being given to valorizing novel and unexploited sources for vegetable oils.
Fig seed represents an interesting atypic source of oil with highly promising lipo-
chemical and biochemical attributes, which can provide many virtues beyond bal-
anced nutrition. Although fig seeds attributes are scarcely documented, oil and
press-cake or meal can be value-added products to be brought to the markets and
used as functional food or for cosmetic applications. Notwithstanding, more
research is required to investigate the properties above further, explore other
research fields such as the varietal effect on phytochemical properties, recovery of
bioactive compounds with different extraction techniques, particularly green ones,
and finally, the stability of FSO according to the above-cited factors. Furthermore,
additional investigations are necessary to explore the effectiveness and technologi-
cal opportunities of using fig oil and cake-press or meal produced as functional
foods or by-products used in formulating new foods. Such research, among others,
might motivate to meet circular economy goals and fulfill the food chain stakehold-
ers’ expectations regarding opportunities to create value for the waste and byprod-
ucts. Therefore, the purpose of this chapter was to draw attention to fig seeds in the
scope of the circular economy by providing a review of the very few available rel-
evant papers on the topic and simply proposing potential valorization pathways, and
to encourage researchers to give more attention to seeds of this outstanding fruit.

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Chapter 15
Fig (Ficus carica) Leaves: Composition
and Functional Properties

Rashida Bashir , Samra Tabassum, Ayoub Rashid, Shafiqur Rehman,

and Ahmad Adnan

Abbreviations

TPC Total phenolic content

TFC Total flavonoid content
TTC Total tannin content
TCC Total carotenoid content
DW Dry weight
FW Fresh weight

1 Extraction of Bioactive Compounds

Maceration is the simplest method of compound extraction. It has been applied to

extract the latex of wild fig “caprifig” using 1 L of n-hexane for 10 days (Lazreg-
Aref et al., 2012); however, simple maceration of latex can be modified with the
help of mechanical shaking. A study conducted on fig latex has shown that macera-
tion with shaking provided better total phenolic content and antioxidants than
ultrasound-assisted extraction at the same reaction conditions especially for the
white genoa variety (Shahinuzzaman et al., 2020).

R. Bashir (*) · S. Tabassum

Division of Science and Technology, University of Education Lahore, Lahore, Pakistan
e-mail: [email protected]
A. Rashid · A. Adnan
Department of Chemistry, GC University Lahore, Lahore, Pakistan
e-mail: [email protected]; [email protected]
S. Rehman
Department of Chemistry, Faculty of Sciences, University of Central Punjab,
Lahore, Pakistan
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 339
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_15
340 R. Bashir et al.

Magnetic stirring is also a standard method used to extract bioactive compounds

from plants; optimum conditions of 115.14 min, 48.6 °C temperature, and 63.4%
acetone have provided high recovery of phenolic compounds and antioxidants
(Bachir Bey et al., 2014). However, Soxhlet extraction requires a higher tempera-
ture than a maceration. Soxhlet extraction has been carried out at 60–80 °C tem-
perature using petroleum ether to recover hepatoprotective compound fraction from
fig leaves (Gond & Khadabadi, 2008). In a study, Soxhlet extraction extracted polar
compounds with antimicrobial activity from fig leaves by using ethanol in a solid/
liquid ratio of 1:7, with 24 h extraction period (Al Askari et al., 2013).
Decoction also requires heat treatment but is confined to water-soluble fractions
only (Hamedi et al., 2015). It has been carried out to recover water-soluble polar
compounds with antibacterial activity; however, did not the antifungal activity was
not reported in the decoction extract of fig leaves (Al Askari et al., 2013). The
hydrodistillation technique also uses water as a solvent but is confined to essential
oils extraction, and GC-MS analysis reported 108 and 121 volatile phytoconstitu-
ents in leaves and fruit, respectively (Li et al., 2012). Therefore, a solid-liquid phase
has been commonly used for compound extraction. However, solid matrix disrup-
tion extraction (sea sand disruption method) of fig leaves provided better reproduc-
ibility (RSD < 5%) and recovery (above 85%) of phenolic acids, flavonols, and
coumarins (Teixeira et al., 2006).
Phenolic acids, flavonols, and coumarins have been efficiently extracted using
ultrasound-assisted extraction using ionic-liquid, which provided better recovery
and reduced extraction time at the optimum conditions of 30 min extraction, 30 °C
temperature, 1:50 solid-liquid ratio, and 1.0 mol/L of imidazolium ionic liquid con-
centration (Hongying, 2015). A tailored deep eutectic solvent system comprising
glycerol, xylitol, and D-(-)-fructose in the molar ratio of 3:3:3 has also been used for
fig extraction; results have indicated that DES assisted UAE and DES assisted MAE
provided a better yield of polyphenolic and furanocoumarin compound as compared
to methanolic and non-tailor DES assisted UAE and MAE. The yield reached
6.582 mg/g, 5.307 mg/g, 16.10 mg/g, 2.575 mg/g, 15.02 mg/g, and 2.575 mg/g for
caffeoylamlic acid, rutin psoralic acid-glucoside, bergapten and, psoralen, respec-
tively, under optimal conditions (Wang et al., 2017).
Supercritical fluid extraction has been used for defatting fig powder before
polysaccharide extraction, using optimum conditions of 45 °C and 30 MPa (Yang
et al., 2009). Microwave-assisted extraction has been used to extract flavonoids
from fig leaves at 70 °C for three times using water and 70% ethanol. Water pro-
vided lesser yield (5.5% w/w) as compared to 70% ethanol (yield = 10% w/w)
(Trifunschi & Ardelean, 2013). Surfactant based microwave-assisted extraction
replaces organic solvent and provides better recovery of bioactive compounds in
less time; polyphenolic compounds and furanocoumarins have been successfully
extracted from fig leaves using this technique with optimal conditions of 40 °C
temperature, 10.27 min, solid/liquid ratio of 1:19.95 (Yu et al., 2020). Enzyme
assisted extraction is a sustainable green method for bioactive compounds; how-
ever, the combination of ultrasound-assisted extraction with enzymatic extraction
gives rise to an innovative technology that reinforces the recovery of polyphenolic
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 341

compounds; Ultrasonic-enzyme assisted two-phase aqueous extraction of fig

leaves has provided higher flavonoid yield of 60.22 mg/g as compared to ultra-
sound-assisted ATPE, enzyme-ATPE, ATPE and Soxhlet extraction (Zhao et al.,
2021). Solid-state fermentation using fungi is an emerging non-conventional tech-
nique reported to enhance the phenolic compound recovery and anti-oxidant activ-
ity up to 5.48 and 98.54 folds from fig processing by-products (Buenrostro-Figueroa
et al., 2017).
The choice of extraction technique and conditions also vary depending upon
target compounds; according to a study, 24 h maceration with ethanol in a 1:50
solid/liquid ratio was used to get extracts for flavonoids, flavonols, ascorbic acid,
proanthocyanidin, and ortho-diphenols estimation however for the extraction of
carotenoids, 5 min ultrasonication using a solvent system of hexane/acetone/metha-
nol was used (Amessis-Ouchemoukh et al., 2017).

2 Phenolic Content and Common Phenolic Acids

Phenolic content is reported to vary among different cultivars of fig. For example, a
study has shown that the total phenolic content ranged between 24–237 mg
GAE/100 g fresh weight for 24 different genotypes and two standard cultivars
(Ercisli et al., 2012). In another study, dark color varieties have shown high TPC,
Fassi variety having the highest TPC of 524 mg GAE/100 g DW, followed by
Noukali, which showed a TPC of 478.6 mg GAE/100 g DW; Snowden also showed
a considerable amount of phenolic compounds showing TPC of 42.52 mg GAE/100 g
DW) (Hssaini et al., 2020a).
The variation in phenolic content is not limited to different varieties only, as a
study has reported that the different parts of the fig had a varied range of phenolic
content, skin being the primary source of phenolic compounds with TPC range from
41.7 to 463 mg GAE/100 g FW followed by fruit 48.6–281 mg GAE/100 g FW
while pulp was experienced low phenolic contents with TPC range of 36.5–100.6 mg
GAE/100 g FW(Solomon et al., 2006).
Extraction temperature and the solvent system have a pronounced effect on the
TPC of fig, a study has shown that 80% ethanol and 80 °C extraction temperature
significantly enhanced the recovery of phenolic compounds (3.7 mg GAE/g dry
basis) from Ficus carica L. cv. Šaraguja (Bucic-Kojic et al., 2011). Methanol extract
of fig latex also has shown a high TPC value of 50.20 mg GAE/g of latex; this value
was approximately five times higher than the TPC of methanol extract of Euphorbia
tirucalli latex (Abdel-Aty et al., 2019). Aqueous methanol extract of whole fruit has
also been reported to have a high TPC value of 31.8% w/w) (Alamgeer et al., 2017).
Extraction technique has pronounced effect on the recovery of phenolic com-
pounds from fig, as a study conducted on eighteen fig varieties has shown that mac-
eration extract of latex gave higher TPC value ranging from 233.1–311.8 μg GAE/
mL as compared to UAE which provided the range TPC range of 224.37–291.98 μg
GAE/mL, however in both types of extraction White Genoa variety yielded the
342 R. Bashir et al.

highest amount of phenolic compounds. In contrast, the Download variety yielded

the lowest TPC, again proving the dependence of TPC on a variety of figs as well
(Shahinuzzaman et al., 2020). Polyphenolic content is reported to vary depending
upon the maturation stages; samples of Dottato variety from June, July, and
September has shown polyphenolic content of 3.9, 4.4, and 11.9 mg chlorogenic
acid equivalent/g of dried material, respectively) (Marrelli et al., 2012).
Plants part, extraction technique, and the solvent system affect the phytochemi-
cal profile (Table 15.1). For example, HPLC analysis of fig fruit has shown the
existence of chronotropic acid, caffeine acid, vanillic acid, syringic acid, gallic acid,
and m-coumaric acid (Alamgeer et al., 2017).
HPLC quantification of phenolic acids of white and dark cultivars of fig fruit has
shown chlorogenic acid to be the most abundant phenolic acid, with the content
ranging from 0.46 to 1.71 mg/100 g FW, followed by gallic acid and synergic acid.
Besides fruit variety and color parameters, phenolic acids were varied in crop culti-
vations from two different months, i.e., July and September (Veberic et al., 2008).
However, another study has not detected gallic acid in fig varieties' pulp (Hssaini
et al., 2021).
HPLC analysis of methanol extracts of fig latex also has not shown the presence
of gallic acid; however, p-protocatechuic acid 8%, chlorogenic acid 59%, vanillic
acid 1%, caffeic acid 3%, sinapic acid 1% were present along with other polyphe-
nolic compounds (Abdel-Aty et al., 2019). When compared among peels and pulp
of two different varieties of fig, protocatechuic acid was present in a higher amount
in the pulp of both varieties (8.11 and 7.45 mg/100 g sample in Calor and Cuello
dama variety, respectively) as compared to peels (6.22 and 3.27 mg/100 g in Calor
and Cuello dama respectively). Chlorogenic acid ranged from 0.92–3.51 mg/100 g
in all samples; however, the pulp of the Calor variety contained chlorogenic acid in
the highest amount of 3.51 mg/100 g (Viuda-Martos et al., 2015). Leaves and twig
of thirteen fig varieties have also been compared for their phenolic acids content,
HPLC analysis identified caffeoylmalic as a predominant phenolic acid in leaves
ranging from 0.79 to 5.97 mg/g DW, 3-O-caffeoylquinic acid (leaves = 1.31–3.54 mg/

Table 15.1 Major phenolic acids in fig

Plant
Major compound Extraction technique part Reference
Chlorogenic acida Shaking Latex Abdel-Aty et al. (2019)
Chlorogenic acidb Ultrasonication Fruit Veberic et al. (2008)
Protocatechuic acida Peels Viuda-Martos et al.
Protocatechuic acida Pulp (2015)
Gallic acida Sonication followed by peels Hssaini et al. (2021)
Chlorogenic acida maceration pulp
Chlorogenic acidb Maceration Leaves Ladhari et al. (2020)
Dihydroxy benzoic Twig
acidb
80% methanol as solvent
a

Methanol as solvent
b
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 343

DW, Twig = 0.97–1.62 mg/g DW), 5-O-caffeoylquinic acid (leaves = 0.34–0.59 mg/g
DW, twig = 0.16–0.36 mg/g DW), dihydroxy benzoic acid(leaves = 1.15–2.15 mg/g
DW, twig = 0.94–2.31 mg/g DW) and caffeoylmalic acid (twig = 0.75–3.66 mg/g
DW) (Ladhari et al., 2020).

3 Flavonoid Content and Common Flavonoids

Flavonoid content estimation of skins, pulp, and fruit has shown a varied range of
TFC depending upon the part of fig understudy, just as TPC. For example, dark-
colored varieties of fig are reported to have higher flavonoid content as compared to
light-colored varieties; however, skins of all varieties have shown higher flavonoid
content (ranged from 2.2 to 45.6 mg catechin/100 g FW) as compared to the pulp
(1.6–5.7 mg CE/100 g FW) and whole fruit (2.1–21.5 mg CE/100 g FW) (Solomon
et al., 2006).
Another study has reported higher TFC in three dark fig varieties (Bouankik,
Azandjar, and Aberkane) with a mean value of 126.55 mg quercetin equivalent/100 g
of dry mass as compared to six light varieties, which showed a mean TFC value of
87.24 mg QE/100 g DM (Bey & Louaileche, 2015). However, another study has
reported higher TFC values for lighter varieties with an average TFC of 37.71 mg
CE/100 g DW, Kadota variety showing the highest TFC value of 83.71 mg CE/100 g
DW while white Adriatic is showing least TFC value of 14.59 mg CE/100 g DW
(Hssaini et al., 2020a). Finally, a study showed the significant influence of steam
explosion pretreatment on flavonoid content of fig leaves extract. Mild and drastic
steam explosion pretreatment resulted in the flavonoid yield of 51.03 mg/g and
44.77 mg/g, respectively. In comparison, untreated leaves showed a flavonoid yield
of 32.73 mg/g after 6 h extraction period (Qin & Chen, 2015).
TFC has shown a positive correlation between extraction temperature and the
percentage of ethanol used for extraction. TFC increased from 0.68 CE/g DB to
2.5 CE/g DB using temperatures from 25 °C to 80 °C; likewise, TFC was observed
from 0.44 mg CE/g DB to 2.5 CE/g DB using 50% to 80% ethanol, respectively
(Bucic-Kojic et al., 2011). The type of extraction solvent also affects the flavonoid
content; ethyl acetate fraction of leaves extract has higher flavonoid content of
83.92 mg/g than water, and petroleum ether extract petroleum-ether gave the least
TFC value of 18.71 mg/g (Li et al., 2021). Using an aqueous methanolic solvent
system has resulted in a high recovery of flavonoids. A study has shown a high TFC
value of 538.20% w/w using aqueous methanol in 70:30 (Alamgeer et al., 2017).
Rutin is a major phytoconstituent of fig (Table 15.2). HPLC analysis of white and
dark cultivars from July and September has identified rutin as a major flavonoid
with a content range from 4.89 to 28.7 mg/100 g FW, followed by catechin, which
showed full content of 4.03 mg/100 g FW(Veberic et al., 2008). Methanol extract of
fig latex also showed a similar trend, with rutin being in the highest percentage of
20%, followed by catechin, which showed a percentage composition of 8%; how-
ever, quercetin and apigenin were not detected in latex at all (Abdel-Aty et al., 2019).
344 R. Bashir et al.

Table 15.2 Major flavonoids compounds in fig

Major Plant
compound Extraction technique part Reference
Rutina Shaking Latex Abdel-Aty et al. (2019)
Rutinb Ultrasonication Fruit Veberic et al. (2008)
Epichtechina Sonication followed by Pulp Hssaini et al. (2021)
Rutina maceration Peels
Rutina Ultrasonication Peels Viuda-Martos et al. (2015)
Catechina Pulp
Rutinb Maceration Leaves Ladhari et al. (2020)
Rutinb Twig
Rutinc Hot plate stirring leaves Belguith-Hadriche et al.
(2017)
a
80% methanol as solvent
b
Methanol as solvent
c
70% ethanol as solvent

Another study conducted on 25 different cultivars has shown epicatechin to be a

significant flavonoid in the pulp of all cultivars, with a mean value of 5.23 μg/g
DW. However, its content was most prominent in the darkest fig cultivar, where it
showed a high content value, up to 54.66 μg/g DW; peels have also shown a consid-
erable amount of epicatechin; other than this, catechin was present in both peels and
pulp; however, quercetin and apigenin were not detected in the pulp (Hssaini et al.,
2021). The absence of quercetin and apigenin accords with the study on methanolic
extract of latex (Abdel-Aty et al., 2019). However, another study has shown the
presence of apigenin along with other flavonoids in peels and pulp of powder co-
product of Calor and Cruella dama fig varieties. Rutin was found to be the most
abundant flavonoid in the peels of both Calor and Cruella dama varieties (15.3 and
14.2 mg/100 g sample, respectively), followed by catechin, which showed the con-
tent value of 6.10 and 14.20 mg/100 g in the peels of Calor and Cruella dama
respectively. Interestingly catechin was the most abundant flavonoid in pulp
(9.53 mg/100 g). Other than these two compounds, luteolin, apigenin, luteolin-7-O-
glucoside, and epicatechin were also present in peels in considerable amounts; how-
ever, luteolin-7-O-glucoside was not detected in the pulp of both varieties
(Viuda-Martos et al., 2015).
UPLC-QTOF-MS analysis has identified 11 flavonoid glycosides in leave extract
obtained by ultrasound-enzyme assisted two-phase aqueous extraction. Only one
compound has already been identified in leave extract; other newly identified com-
pounds include quercetin glycosides, apigenin, kaempferol, luteolin, and napthohy-
droquinone (Zhao et al., 2021). Study of leaves and twig of 13 fig varieties has
identified three flavonoid glycosides including quercetin-3-O-glucoside
(leaves = 4.73–7.51 mg/g DW, twing = 2.49–6.45 mg/g DW), quercetin-3-O-
rutinoside, (leaves = 12.43-22.70 mg/g DW, twig = 8.14–17.23 mg/g DW) and
kaempferol-3-O-glucoside (leaves = 5.36–12.45 mg/g DW, twig = 637–10.23 mg/g
DW) (Ladhari et al., 2020). RP-LC-DAD-QTOF-MS analysis of leaves extract of
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 345

two Tunisian cultivars has identified thirteen flavonoids molecules of different sub-
classes, including flavonols (rutin and isoquercetin), flavones (luteolin C-hexoside
C-pentoside I, luteolin C-hexoside C-pentoxide II, Apigenin C-hexoside C-pentoside
I, Apigenin C-hexoside C-pentoxide II, vitexin, luteolin) and isoflavones (cajanin,
prenylhydroxygenistein I, prenylhydroxygenistein II, prenylgenistein I, prenylge-
nistein II) (Belguith-Hadriche et al., 2017).

4 Anthocyanin

Anthocyanin content in fig also varied between different varieties as a study showed,
anthocyanin content varied in the range of 0–42 μ cyanidine-3-rutinoside/g FW in
different genotypes and cultivars of figs. It also found that the presence of anthocya-
nins influenced the color of figs, as black figs showed higher anthocyanin content
than purple or green colored figs (Ercisli et al., 2012). Another study carried out on
nine different fig varieties has also indicated higher anthocyanin content in dark
varieties with the highest anthocyanin content in the Bouankik variety, which
showed anthocyanin content of 20.78 mg quercetin-3-glucoside equivalent/100 g
DM followed by Aberkane (18.73 mg Q3GE/100 g DM) and Azandjar (17.18 mg
Q3GE/100 g DM), this content range is much higher than that was observed for
light varieties which showed the mean content of 4.9 mg Q3GE/100 g DM (Bey &
Louaileche, 2015). Hssaini reported similar results, where black-colored varieties
Noukali, Fassi, and Ghoudan showed high anthocyanin content of 23.77, 14.88
11.16 mg Cy-3-rutinoside equivalent/100 g DW, respectively. Anthocyanin content
comparison in skin and pulp of five varieties; Colar, Cuello de Dama (green and
dark purple), Granilla, and Bursa, have shown anthocyanin content to be several
times higher in the skin (range of 32–97 μg/g FW) with highest anthocyanin content
in the skin of Bursa siyahi variety of fig as compared to a pulp (1.5–15 μg/g FW)
where the pulp of Colar variety showed highest AC (Dueñas et al., 2008). This study
is followed by the findings of Solomon et al. (2006), where the skin extract samples
of fig showed the highest range of anthocyanin content (0.7–27.3 mg cyanidin-3-
glucoside/100 g), followed by fruit extract(0.3–10.9 mg cyanidin-3-glucoside/100 g).
Another study has shown the highest AC in peels of purple variety, 196 mg
Cy-3-G/100 g DW; however, drying of fig can cause loss of anthocyanin content;
according to a study, the anthocyanin content decreased up to 83 and 98% upon dry-
ing in purple and yellow fig variety respectively. AC ranged from 0.8 to 5.9 mg
Cy-3-G/100 g DW in different fig parts of the yellow variety, which decreased to
0.1 mg Cy-3-G/100 g DW upon drying, while in the purple variety, the AC ranged
from 38 mg to 196 Cy-3-G/100 g DW that declined to 14.5 mg Cy-3-G/100 g DW
(Kamiloglu & Capanoglu, 2015).
HPLC-DAD-MS analysis of different fig varieties comprehensively profiled the
anthocyanin composition of fig. Primary aglycon moiety was reported to be cyani-
din, and glycon substituents were found to be rutinose and glucose. Cyanidin-3-
rutinoside was the most abundant anthocyanin in both skin and pulp, constituting
346 R. Bashir et al.

48–81% of total anthocyanins in the skin and 68–79% of total anthocyanins in the
pulp. Cyanidin-3-glucoside was the second prominent anthocyanin after cyanidin-3-
rutinoside (Dueñas et al., 2008). These results are per a study conducted on twenty-
five fig varieties by Hssaini et al. (2021), which also identified cyanidin-3-rutinoside
as the most abundant anthocyanin in both plant parts, i.e., peel and pulp; however,
higher content of cyanidin-3-rutinoside (77.97 μg/g DW) was present in peels of as
compared to the pulp (9.01 μg/g DW) other than cyanidin-3-rutinoside, cyanidin-3,5-
diglcoside was also present in high amount in peels (75.90 μg/g DW). It also found
that the darkest colored cultivar provided the most promising anthocyanin profile
with high content values of anthocyanin compounds, most prominently cyanidin-3-
rutinoside and cyanidin-3, 5-diglcoside with the content value of 478.6 μg/g DW
and 494.0 μg/g DW, respectively. Another study conducted on powder co-product of
Calor and Cuello dama fig varieties has shown higher cynaidin-3-rutinoside and
cyanidin-3-glucoside (2.01 and 1.33 mg/100 g respectively) in peels of Calor vari-
ety as compared to Cuello dama variety (0.43 and 0.12 mg/100 g respectively),
however, both of these significant anthocyanidins were not detected in the pulp of
both varieties (Viuda-Martos et al., 2015). Cyanidin-3-rutinoside and cyanidin-3-
glucoside accumulation in the skin highly impact the fruit color; the accumulation
of these compounds could be enhanced by the treatment of exogenous abscisic acid
and ethephon during fruit ripening. Cyanidin-3-O-glucoside increased up to 31.66
and 12.35 μg/g FW after 72 h of exogenous ABA and ethephon treatment, respec-
tively, compared to control (5.47 μg/g FW). Cyanidin-3-O-rutinoside reached the
level of 10.39 μg/g FW after 72 h of exogenous-ABA treatment, while ethephon
treatment showed cyaniding-3-O-rutinoside content of 4.84 μg/g FW as compared
to control (2.45 μg/g FW) (Lama et al., 2020).

5 Proanthocyanidins

Proanthocyanidins are also a prominent class of bioactive compounds present in

figs, and a research study has shown high proanthocyanidin content of 40 mg/g in a
fig variety of Spain (Vallejo et al., 2012). Furthermore, the proanthocyanidin con-
tent of fig (Ficus carica) was reported to be even higher (496.6 mg CE/100 g DW)
than Ceratonia siliqua (140 mg CE/100 g DW) and Quercus ilex (254.5 mg
CE/100 g DW) (Amessis-Ouchemoukh et al., 2017).
Proanthocynidin content varies between different colored varieties, just as antho-
cyanins; when dark and light varieties were compared, Aberkane (a dark variety
showed the highest proanthocyanidin content of (179.8 mg cyanidin equiva-
lent/100 g dry mass); however, Abiarous (a light-colored variety) showed lowest
proanthocyanidin content of 103.3 mg cyanidin equivalent/100 g DM (Bey &
Louaileche, 2015). However, another study has shown that light color varieties con-
tained higher proanthocyanidin content than dark varieties; Cuello Dama Blanca
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 347

and Snowden varieties showed the highest proanthocyanin content of 6.64 cyanidin
equivalent/100 g DW, respectively (Hssaini et al., 2020a).
Like anthocyanins, drying of figs is also reported to impact proanthocyanidin
content. The purple fig variety was found to have higher proanthocyanidin content
than the yellow variety, but drying decreased the content up to 74% in purple fig.
Conversely, an increase of proanthocyanin content up to 71% was witnessed in yel-
low varieties upon drying. In yellow varieties, the content in different parts varied
from 6.6 to 7.2 mg CyE/100 g DW, which increased to 12 CyE/100 g DW in dried
fig. Similarly, in the purple fig, the content varied from 34 to 220 CyE/100 g DW,
which upon drying declined to 16 CyE/100 g DW (Kamiloglu & Capanoglu, 2015).

6 Coumarins

Psoralen and bergapten are major coumarins that have been detected in the aqueous
methanolic extract of fig leaves (Teixeira et al., 2006). An HPLC-UV analysis con-
ducted on leaves, pulp, and peels of fig also has shown the highest psoralen and
bergapten content in leaves (67.8 μg/0.1 g sample and 20.2 μg/0.1 g sample, respec-
tively), followed by pulp (psoralen, 5.56 and bergapten, 2.49) however peels showed
the lowest content (psoralen, 2.59 μg/0.1 g sample and bergapten 1.99 μg/0.1 g
sample) (Hongying, 2015).
Bark and wood of fig plant have reported high levels of psoralen and bergapten,
and psoralen ranged from 146.6–1110.3 μg/g DW in bark and 395.7–1671.8 μg/g
DW in wood, while bergapten was lower in concentration as compared to psoralen,
ranging from 144.2 to 718.6 μg/g DW in wood while 114.3 to 524.0 μg/g DW in the
bark.(Rouaiguia-Bouakkaz et al., 2013) Other than psoralen and bergapten,
8-methoxypsoralen, angelicin, rutaretin, pimpinellin, and seselin have also been
detected in the Dottato variety of fig. Psoralen was predominant coumarin, just as in
other studies (23% peak area in GC-FID); however, it was followed by seselin in its
content value (19.50% peak area) rather than bergapten (15.20% peak area).
Extraction solvent affected the recovery of coumarins as n-hexane provided better
recovery than dichloromethane (Marrelli et al., 2012).
RP-LC-DAD-QTOF-MS of leaves extract of Temri, and Tounsi varieties of fig
have shown three hydroxycoumarins including Phellodenol A(hydrated 4′,5′-dihy-
dropsoralen), Murrayacarpin B/dihydrated bergapten) and prenyl-7-
hydroxycoumarin and two furanocoumarins hydroxylpsoralen hexoside and prenyl
methoxypsoralen (Belguith-Hadriche et al., 2017). GC-MS profile of n-hexane
extract of latex of caprifig (wild fig variety) has shown coumarins to be the major
constituents of extracts; fourteen coumarins were identified, constituting 90.56% of
total identified compounds (Lazreg-Aref et al., 2012).
348 R. Bashir et al.

7 Tannins

Tannins are complex bioactive compounds derived from phenolic acids; a study car-
ried out on three fig varieties has shown higher total tannin content in green varieties
(Bidhi) as compared to purple (Hamri) and black varieties (Kohli). Bidhi peels con-
tained the highest full tannin content of 75.98 mg tannic acid equivalent/g FW, fol-
lowed by pulp (65.87 mg TAE/g FW), while whole fruit represented TTC of
60.35 mg TAE/g FW. Peels of the Kohli variety also contained more tannins than
pulp and whole fruit; however, the Hamri variety showed the highest concentration
of tannins in the entire fruit (Harzallah et al., 2016).
Studies have shown that Taamriout and Azendjar varieties are rich in condensed
tannins compared to hydrolyzable tannins. Methanol extracts have shown the high-
est TTC ranging from 160 to 194 mg GAE/100 g DW. In comparison, petroleum
ether extract of Azendjar has shown the most negligible TTC value (below 100 mg
GAE/100 g DW) (Debib et al., 2014). In another study carried out on the estimation
of condensed tannin, peels of Bakkor Khal and pulp of Zarrouk showed the highest
condensed tannin content of 18.4 and 4.4 μg catechol equivalent/g FW, respectively;
however, the Bither variety showed the lowest CT content range of 0.388–0.607 μg
CE/g FW) (Mahmoudi et al., 2018).

8 Carotenoids

Carotenoid's presence is also identified in fig fruit; a study has identified three carot-
enoids in fig fruit, including β-carotene, α-carotene derivative, and β-cryptoxanthin.
β-carotene was present in the highest amount of 4.32 μg/100 g DE followed by
β-cryptoxanthin (2.14 μg/100 g DE) (Amessis-Ouchemoukh et al., 2017).
Yellow fig varieties have reported four carotenoids: lutein, zaexanthin,
β-cryptoxanthin, and β-carotene. Upon comparing fresh samples, the Sarilop vari-
ety showed higher total carotenoid content (29.6 μg lutein equivalent/g DW) as
compared to the Sarizeybek variety (23.7 μg LE/g DW); however, lutein content
was higher in Sarizeybek (7.15 μg/g DW) than Sarilop variety (6.14 μg/g DW). It
also proved that fig drying negatively affects the total carotenoid content as TCC
decreased up to 80% upon seven-day drying (Yemis et al., 2012).
A study reported the effect of storage duration on total carotenoid content; it was
shown that Peels of the Dottato variety had higher total carotenoid content
(17.8 μg/g) as compared to Melanzana at five day storage time. On the other hand,
Melanzana showed an evident decline in leaves TCC over the storage period of
10 days. However, carotenoids in the pulp of both varieties were stable upon storage
even for 10 days showing a total carotenoid content of approximately 3.0 μg/g
(Allegra et al., 2017).
Carotenoid content tends to be different in hand-pollinated and parthenocarpic
figs, which decreases during the maturation of fig. In pollinated figs, the carotenoid
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 349

content decreased from 18.39 to 7.71 μg/g FW, while in parthenocarpic figs, the
carotenoid accumulation reduced from 16.62 to 6.64 μg/g FW. Individual carot-
enoids were found to be lutein, b-carotene, zeaxanthin, and violaxanthin, lutein
being the major carotenoid with the content value of 9.63 μg/g FW at 0 weeks of
pollination, which increased to 12.58 μg/g FW after 1 week and decreased to
4.43 ug/g FW after 5-week maturation (Nawade et al., 2020).

9 Terpenes and Other Volatile Compounds

GC-MS analysis of fig leaves essential oil has shown the presence of 66 volatile
compounds, among which the oxygenated compounds were most the abundant,
constituting the 46.3% of volatile compounds composition, followed by aliphatic
hydrocarbons (28.2%), fatty acids (9.9%) and sesquiterpenes (3.1%) however
monoterpenes were least abundant exhibiting 0.6% of the total composition. The
prominent compounds belonging to these classes included oxygenated compounds;
(Z)-3-hexeny benzoate, (E)-2-hexenal, phytol and n-nonanal, aliphatic hydrocar-
bons; n-tetracosane and n-docosane, sesquiterpene; (E, E)-α-farnesene and cis-
muurola-4(15),5-diene, monoterpene; α-pinene (only monoterpene in understudy
oil), fatty acid; n-hexadecanoic acid (Ayoub et al., 2010).
Fig fruits samples from different localities have been studied for their flavoring
agents, and primary flavoring agents included p-cymene, decanol, carvacrol, trico-
sane, tetracosane, pentacosane, heptacosane, octacosane, and nonacosane. It also
found that geographical location influences the volatile composition of figs (Darjazi
& Larijani, 2012).
Solid-phase micro extraction-GC-MS has been used for identification and
quantification of volatile compounds from fructus (fruit) and folium (leaves) of
fig; fig fructus has shown the presence of only one volatile compound, 2,-butane-
diol by using the SPME technique; however, SPME-GC-MS has demonstrated the
presence of tetramethyl-decane, trimethyl-undecane, octadecane, carvacrol,
caryophyllene oxide, and β-caryophyllene. SPME detection of folium has shown
3, 5-octadiene-2-on, trimethyl-methylene-vinibicyclononane, hexanal, undecane,
dimethyl- undecane, and ethyl-cyclohexanol; however, folium steam distillate
detected by SPME-GC-MS has identified methyl-undecane, dimethyl-undecane,
nanodecane, β-caryophyllene, caryophyllene-oxide and a trace amount of
β-ionone (Ficsor et al., 2012).
Hydrodistillate of fig fruit of six different cultivars (Bither Abiadh, Bidi, Bither
Kholi, Himri, Kholi, and Tchich Asal) have shown higher cedrol (43.8%) and
α-pinene (9.3%) as compared to leave distillate which showed the percentage of
38.9 and 6.1% for cedrol and α-pinene respectively, however many oxide (24.8%)
and abietatriene (11.8%) was higher in leaves than fruits (12.9 and 8.1% of manoyl
oxide and abietatriene respectively). Other than these compounds, leaves hydrodis-
tillate also showed the presence of α-terpineol acetate (21.7%), γ-muurolene (7.4%),
350 R. Bashir et al.

pentadecanal (5.2%), and nonadecanal (2.3%) however, fruits showed the presence
of α-terpinyl acetate (22.5%), tans-calamenene (3.9%) and n-heneicosane (3.5%)
(Soltana et al., 2017).
GC-MS-Olfactometric analysis of fig leaves pulp has identified almost 58 aroma
compounds collectively in peels and pulp of Sarilop and Bursa Siyahi varieties of
fig. That composition mainly comprised this composition of terpenes followed by
ester and alcohols. The highest terpenes content was seen in peels of Sarilop variety
that showed the content value of 7691 μg/kg FW; however, among 21 individually
detected terpenes DL-limonene showed the highest content, ranging from 441 to
3320 μg/kg FW, followed by β-caryophyllene that delivered the content range of
556–2757 μg/kg FW. Other terpenes include D-3-carene, p-xylene, γ-terpinene,
o-cymene, citronellal, α-cubebene, α-copaene, linalool, α-guaiene, humulene, ger-
macrene, alloaromadendrene, β-bisbolene, α-farnesene, (Z)-α-bisabolene, caryoph-
yllene oxide, and carvacrol (Sertkaya et al., 2021).
GC-MS analysis of n-hexane extract of caprifig also identified eight terpene
compounds constituting 4.81% of the total identified compounds. Among these ter-
penes, sesquiterpenes were most abundant, with a percentage composition of 2.83%
(germacrene-D, delta-cadiene, 2,6,10-Dodecatrien-1-ol), followed by two oxygen-
ated triterpenes (0.74%) and two triterpenes (0.46%); however, bornanone-3 was
the only identified monoterpene present in the extract (Lazreg-Aref et al., 2012).
β-bourbonene, α-guaene, α-muurolene, τ-muurolene, τ-cadinene, α-cubenene,
β-cubenene, menthol, 1-penten-3-ol, β-pinene, hydroxyl caryophyllene, and euca-
lyptol are terpenoids derived from leaves, fruit, and bark of fig which have been
studied with the help of molecular docking and dynamic simulation for understand-
ing the anticancer activity of the plant. Among all mentioned compounds,
β-bourbonene, a sesquiterpenoid, has shown promising results (Gurung et al., 2021).

10 Fatty Acids and Phytosterols

Latex of fig has been reported as a source of fatty acids. Saturated fatty acids majorly
contribute to latex's total fatty acid profile, constituting approx. 86% of it, arachidic
being the most abundant saturated fatty acid (44.1%), followed by palmitic acid
(21.4%) and behenic acid (13.1%) (Oliveira et al., 2010).
GC analysis of seeds oil of four fig varieties has shown a high amount of linole-
nic (38.4–43.5%) followed by linoleic acid (28.9–34.5%), both of which are poly-
unsaturated fatty acids. Among saturated fatty acids, palmitic acid was predominant,
showing a content range from 8.54 to 9.05%, followed by stearic acid, which
showed a content range from 2.59 to 3.3% (Hssaini et al., 2020b). Not only seeds
and latex fig leaves are also found to be a source of fatty acids; however, the content
percentage is low (18% of total ion current, identified by GC-MS), and leaves
extracts analysis has shown the presence of myristic acid, palmitic acid, margaric
acid, linoleic acid, α-linolenic acid, stearic acid, arachidic acid, behenic acid, ligno-
ceric acid, and cerotic acid. Palmitic and stearic acids were predominant among
15 Fig (Ficus carica) Leaves: Composition and Functional Properties 351

saturated acids, showing content from 4% to 14% and 1% to 4% of TIC, respec-

tively. In comparison, primary polyunsaturated acids were found to be α-linolenic
acid (5–13.8% of TIC) and linoleic acid (1.8–3.9% of TIC) (Ivanov et al., 2018).
The fig fruit has also shown 20 fatty acids collected during June, July, and
September, including methyl ester of hexanoic acid, the ethyl ester of (E)-2-
heptenoic acid, caprylic acid ethyl ester, decanoic acid, pentadecanoic acid, ethyl
ester, palmitic acid methyl ester, palmitic acid, myristic acid, palmitoleic acid ethyl
ester, palmitic acid ethyl ester, 14-hexadecanoic acid methyl ester, pentadecanoic
acid, margaric acid ethyl ester, linoleic acid methyl ester, stearic acid methyl ester,
stearic acid, oleic acid ethyl ester, linoleic acid ethyl ester and stearic acid ethyl
ester (Marrelli et al., 2012).
Phytosterols are also reported in fig latex extracts. GC-ITMS and HPLC analysis
of fig latex has identified seven phytosterols, including b-amyrin, a-amyrin,
b-sitosterol, lupeol, lupeol acetate, betulol, and lanosterol. Among these identified
compounds, b-sitosterol was the most abundant phytosterol representing 54% of the
total estimated phytosterols. In contrast, the least abundant phytosterol was
a-amyrin, constituting only 0.4% of total phytosterols (Oliveira et al., 2010). GC-MS
analysis of leaves extracts of fig has also identified the presence of phytosterols
(18% of total ion current), including stigmasterol, β-sitosterol, lanosterol, and
cycloartenol. Among all reported phytosterols, β-sitosterol was found to be the most
abundant (11–14% of TIC), followed by cycloatenol acetate (5–9% of TIC) (Ivanov
et al., 2018). Fig achenes oils have been investigating the presence of phytosterols;
according to a study, fig achene oil of the kholi variety has shown a high phytosterol
content of 1061 mg/100 g, where stigmastadienol was identified as the most abun-
dant phytosterol with the content value of 73.7% (Soltana et al., 2016).

11 Vitamins and Other Compounds

α-Tocopherol is a kind of vitamin E reported in fig by many studies; according to a

study, n-hexane leaves extract of fig contains α-tocopherol in a high amount of
57 mg/g dry leaves (Konyalιoğlu et al., 2005). Ascorbic has also been identified in
fig leaves extract in 21.7 to 22.4 mg ascorbic acid/100 g DW. In the same study,
vitamin E content showed variation according to the size of the leaves; large leaves
showed slightly higher vitamin E content (1.9 mg vitamin E/100 g DW) as com-
pared to small leaves (1.8 mg vitamin E/100 g DW) (Ghazi et al., 2012).
Turkish variety of figs has shown a diverse range of vitamins; however, the con-
tent varied in different sections of figs. γ-Tocopherol, ẟ-tocopherol, vitamin D2,
vitamin D3, α-tocopherol, α-tocopherol acetate (acetate), and vitamin K1were iden-
tified in fruit, peel, red and white sections of fig. γ-tocopherol was predominant
vitamin showing the highest content in the skin followed by a white section of fig
(Guvenc et al., 2009).
352 R. Bashir et al.

Vitamin figs have also shown the presence of crude alkaloids (9.6 g/100 g DM)
and saponins (0.59 g/100 g DM) in the edible portion; however, not much data is
available in this instance (Soni et al., 2014).

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Chapter 16
Fig (Ficus carica) Seed Oil

Emi Grace Mary Gowshika Rajendran

Abbreviations

FAO Food and Agricultural Organization

SFA Saturated Fatty Acid
PUFA Polyunsaturated Fatty Acid
MUFA Monounsaturated Fatty Acid
LDL Low Density Lipoprotein
GAE Gallic Acid Equivalent
TH Taamriwthe
AZ Azegzaw
AV Averkane
GC/MS Gas Chromatography/Mass Spectrometry

1 Fig Seeds and Its Oil

More than 750 varieties of fig fruits, which come under the Moraceae family, were
identified (Guvenc 2009). The edible fruit species among these varieties were Ficus
carica L. The forms of consumption of figs can be dried, raw, canned or through
other preserved forms (Mawa et al., 2013). The countries (Fig. 16.1) with hot and
dry summers and very mild winters are the notable producers of this fruit, including
Egypt, Spain, Brazil, Turkey, California, Morocco, Italy, Greece, and other coun-
tries. It is reported that 30% of the world’s fig production comes from Turkey
(Nakilcioglu & Hisil, 2013; Mehta et al., 2014).
The fig seeds are the most attractive part, providing effective health benefits and
nutritional value. A jelly-like flesh is bound within a seed mass, which is contained

E. G. M. G. Rajendran (*)
Women’s Christian College, Chennai, India
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 357
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_16
358 E. G. M. G. Rajendran

Fig. 16.1 Distribution of fig production by Country (2017). (Source: FAO, 2020)

in the white inner ring on the interior portion of the fig (Badgujar et al., 2014). The
seed number varies between 30–1600 per fruit and is minute, small, medium, or
large (Lansky et al., 2008; Badgujar et al., 2014). The fig seeds are hollow unless
pollinated, and numerous edible seeds are found. In addition, the seeds provide the
characteristic nutty flavor of the dried figs, which are now used as lubricants and
edible oils (Badgujar et al., 2014).
The nutritive value of this fruit makes it an important constituent in the
Mediterranean diet, which includes energy, vitamins, dietary fiber, minerals, anti-
oxidant compounds (phenolics), and amino acids (Oliveira et al., 2009; Bucic-kojic
et al., 2011; Amessis-Ouchemoukh et al., 2017). In addition, many usages in thera-
peutics from the roots, fruits, and leaves of the fig are widely known, especially for
respiratory (cough, sore throat, bronchial issues), anti-inflammatory, cardiovascular,
gastrointestinal disorders (indigestion, diarrhea, colic, loss of appetite) and anti-
spasmodic remedies (Mawa et al., 2013; Amessis-Ouchemoukh et al., 2017).
The studies on the chemical composition (Table 16.1) of figs (Ficus carica) have
been extensively done due to its traditional curative properties and its reputation in
various fields (Penelope, 1997; Duke et al., 2002; Lansky et al., 2008; Joseph & Raj,
2011; Badgujar et al., 2014; Rahmani & Aldebasi, 2017). However, using fig seeds
as a beneficial food source is not entirely possible due to their small size and indi-
gestibility, leading to elimination via feces.
The fig seeds contained 18 fatty acids, and statistically significant differences
were found between different locality seeds (Table 16.2). The fatty acid composi-
tion of fig seeds was comparatively less than fresh whole figs (Guvenc 2009).
16 Fig (Ficus carica) Seed Oil 359

Table 16.1 Chemical Composition of Stem, Leaves, Latex and Seeds of Ficus carica
Stem Leaves Latex Seeds
Campesterol T Moisture (%) 67.6 Caoutchouc 2.4% Oleic 18.9%
acid
Hentriacontanol T Protein (%) 4.3 Resin T Linoleic 33.7%
Stigmasterol T Fat (%) 1.7 Albumin T Linolenic 32.9%
Euphorbol T Crude fiber 4.7% Cerin T Palmitic 5.23%
Hexacosanate T Ash (%) 5.3 Sugar T Stearic 2.18%
Ingenol T N-free extract 16.4% Malic acid T Arachidic 1.05%
Taraxerone T Pentosans 3.6% Rennin T
Carotene T Proteolytic enzymes T
Bergaptene T Diastase T
Stigmasterol T Esterase T
Sitosterol T Lipase T
Rutin T Catalase T
Sapogenin T Peroxidase T
Calotropenyl T
Lepeolacetate T
Oleanolic T
T present in trace amounts
Adapted from Joseph and Raj (2011)

The γ-linolenic acid and linoleic acid were higher in fig seeds (41.8%) than in fresh
figs. (1.46%). The Germencik location fig seeds contained the highest amount of
PUFA and the lowest SFA. Fig seed oil is nutrient-rich, and a healthier source of oil
as the essential fatty acids contribute to human health.
Higher contents of α-linolenic and linoleic acids, 32%–50% and 20%–35%,
respectively, are found in fig seed oil. These PUFA acids are not synthesized in
mammals but should be directly consumed through the diet, thereby classifying
them as essential fatty acids (Field, 2003). Omega-3 fatty acids enhance ocular
health and reduce the risk of age-related macular degeneration and cardiovascular
diseases through their anti-inflammatory properties. In addition, fig seed oil reduces
C-reactive protein levels through endothelial functions and protects the cell mem-
branes from free radical damage through its vitamin E content (Richer et al, 2016).
New and under-utilized sources having adequate PUFA content are being dis-
covered and formulated due to the rise in nutritional interest for healthy oils. Fig
seed oil plays a preventive role in reducing LDL and total cholesterol levels, fur-
ther preventing cardiovascular diseases (Ajayi & Ajayi, 2009). The increase in
unsaturated fatty acid content terms fig seed oil as an important source of healthy
essential oils. The oil could be obtained by processing figs and separating the seeds
from the pulp. This could be used to formulate many dietary supplements and food
products.
360 E. G. M. G. Rajendran

Table 16.2 Fatty acid compositions (%) of Ficus carica seeds

Location
Fatty acid Common name Turkey Incirliova Germencik Nazilli
C16:0 Palmitic acid 5–9% 7.4% 3.58% 7.0%
C16:1 Palmitoleic acid – 0.06% 0.06% 0.05%
C17:0 Heptadecanoic acid – 0.06% 0.06% 0.06%
C18:0 Stearic acid 2–4% 3.38% 3.73% 2.97%
C18:1 (n-9) Oleic acid 14–24% 16.8% 17.7% 16.9%
C18:2 (n-6) Linoleic acid 20–35% 32.8% 37.9% 31.8%
C18:3 (n-3) α-Linolenic acid 32–50% 0.30% 0.25% 0.30%
C18:3 (n-6) γ-Linolenic acid 0.17% 37.8% 41.8% 39.1%
C20:0 Arachidic acid Max. 0.7% 0.02% 0.03% 0.02%
C20:1 (n-9) Eichosenoic acid 0.5% 0.09% 0.04% 0.09%
C22:0 Behenic acid Max. 0.5%
C21:0 Heneicosanoic acid – 0.04% 0.08% 0.04%
C20:3n6 Eicosatrienoic acid – 0.09% 0.04% 0.09%
C24:0 Lignoceric acid – 0.03% 0.04% 0.03%
C20:5n3 Eicosapentaenoic acid – 0.39% 0.24% 0.11%
SFAs – 10.9% 7.10% 10.2%
MUFAs – 17.2% 17.0% 17.4%
PUFAs – 71.8% 75.8% 72.2%
SFAs saturated fatty acids; MUFAs monounsaturated fatty acids; PUFAs polyunsaturated fatty acids
Adapted from Guven et al. (2019) and Tas (2018)

2 Extraction and Processing of Fig Oil

The ripe figs were plucked from the branches, peeled manually, and dried using a
grill. After drying the figs, they are cut into small pieces (5 mm), rinsed, precipi-
tated, and dried. The dried fig pulp is separated from the seeds and dried naturally.
These seeds are converted into pure seed oil through an extrusion process from the
cold-extrusion oil processing machine with a temperature below 25 °C. The pure
Ficus carica seed oil (Fig. 16.2) is preserved at −18 °C for further processing and
formulation into different food products and supplements.
From the estimation of sterol composition (Table 16.3), the highest composition
of sterol present in the fig seed oil was β-sitosterol (66.4%). On the contrary, the
lowest was clerosterol (0.68%), and the total sterol content in the fig seed oil was
6516 mg/kg.
16 Fig (Ficus carica) Seed Oil 361

Pure seed oil

extruded is
preserved at -18°C

Extrusion process
Ripe figs- plucked,
using cold extrusion
peeled manually,
oil processing
dried on grill
machine at 25°C

Dried fig pulp is

Aer drying, it is cut
separated from
into small pieces
seeds and seeds are
(5mm) size
dried naturally

The cut pieces are

rinsed, precipitated
and dried

Fig. 16.2 Processing and Extraction of fig (Ficus carica) seeds into pure fig oil. (Adapted from
Baygeldi et al. 2021)

Table 16.3 Sterol composition in Fig (Ficus carica) seed oil

Sterols Sterol Composition %

Beta-sitosterol 66.4
Delta-5-avenasterol 20.1
Delta-7-avenasterol 5.07
Campesterol 2.99
Sigmasterol 2.18
Delta-7-sigmasterol 1.28
Sitosterol 1.18
Clerosterol 0.68
Total Sterol content 6516 mg/kg
Reproduced from Guvenc (2009)
362 E. G. M. G. Rajendran

3 Minor Bioactive Compounds

The fat-soluble vitamin that acts as a potent antioxidant reducing agent in vegetable
oils, known to occur through the chain-breaking of saturated fats, is known as
tocopherol (Shin et al., 2009). However, some minor bioactive compounds that exist
predominantly in fig seed oil include phytosterols, phenolic compounds, organic
acids, and anthocyanins that enhance antioxidant activity (Mawa, Husain &
Jantan, 2013).
From the two location variation of Ficus carica seed oil in Turkey (Table 16.4),
it was found that gamma-tocopherol content was the highest (391.8 mg/100 g) and
(4267 ppm). There was a variation between the delta and alpha-tocopherol compo-
sition, where the delta-tocopherol was higher in the Aydin variety (7.65 mg/100 g)
and comparatively less than Istanbul variety (147 ppm), and alpha-tocopherol was
higher in the Istanbul variety (157 ppm) comparatively more than Aydin variety
(4.6 mg/100 g). Both the varieties did not detect the presence of beta tocopherol. It
was found that the content of gamma-tocopherol was present due to the extraction
of oil from the seeds.
The phenolic compounds were extracted using slight modifications from the
method adopted in the study by Tsimidou et al. (1992). First, in 10 ml of hexane, 2 g
of oil sample was dissolved. Next, this mixture was added to 4 ml of 60% methanol.
The mixture was kept in darkness for 2 h, during which it was subjected to agitation
and filtered using Whatman No.1 filter paper. The second mixture was extracted
using the same method and under the same conditions. First, the two filtrates
obtained were mixed. Then, using a rotary evaporator, the mixture was concentrated
using a vacuum and was reconstituted using 10 ml of pure methanol. This was
stored at 20 °C.
The total phenolic content was determined based on the Folin-Ciocalteu colori-
metric method (Favati et al., 1994). This reaction mixture comprises 250 μL of
Folin-Ciocalteu reagent, 50 μL of extract, 750 μL of sodium carbonate (7%), and
3 ml of distilled water. A 950 μL of distilled water was added after 8 min of stirring
under ambient temperature. The absorbance was measured against a 760 nm blank
after 1 h incubation in the dark. The standard used for the calibration curve was gal-
lic acid. The analysis was carried out in triplicates and was expressed as mg equiva-
lent of Gallic acid (GAE) per 100 g of oil. Therefore, fig seed oil‘s total phenolic
content was 79.5 mg (GAE)/ 100 g of the oil (Guvenc 2009).

Table 16.4 Tocopherol content in fig (Ficus carica) seed oil

Aydin, Turkey Istanbul, Turkey

Type of tocopherol mg/100 g ppm
Gamma-tocopherol 391.8 4267
Delta-tocopherol 7.65 147
Alpha-tocopherol 4.6 157
Beta-tocopherol ND ND
ND not detected, ppm parts per million
Adapted from Guvenc (2009) and Baygeldi et al. (2021)
16 Fig (Ficus carica) Seed Oil 363

4 Antioxidant Activity and Other Health-Promoting Effects

In a study, six different varieties of figs varying in color (yellow, black, red, and
green) were studied for the antioxidant properties present in the oil when seeds are
extruded from dried fig pulp. They were analyzed for their anthocyanin profile, total
polyphenols, antioxidant capacity, and flavonoids. It was reported that 28% and
36% of the total antioxidant capacity was contributed by anthocyanin fraction in red
Brown-Turkey and dark-colored mission varieties of fig oils. In addition, 92% of the
antioxidant capacity in the anthocyanin fraction was contributed by cyaniding-3-O-
rutinoside (C3R). The mission variety fig oils exhibited the highest antioxidant
capacity with the highest flavonoids, polyphenols, and anthocyanin levels (Joseph
& Raj, 2011). Studies evaluating the numerous health benefits of Fig (Ficus carica)
seed oil have been tabulated (Table 16.5).
Fig is a small deciduous plant with immense therapeutic effects owing mostly to
its antioxidant capacity. However, more animal and human clinical trials are required
to confirm and confer other health benefits.

5 Volatile Compounds’ Contribution

to Organoleptic Properties

From a study conducted by Zidi et al. (2021), the volatile compounds identified
from three cultivars of figs through which the oil was extruded from their seeds had
identified almost 29 compounds (Table 16.6) during three different ripening stages

Table 16.5 Therapeutic roles of fig (Ficus carica) seed oil from clinical studies
Therapeutic Role The outcome of the Study References
Anti-oxidative Effective scavenging activity was observed in fig oils Yang et al.
(2009)
Anti-cancer Stomach cancer cell line growth was inhibited Hashemi et al.
(2011)
Anti-cancer Inhibitory effects on proliferation of cancer cell lines by Rubnov et al.
6-O-acyl-beta-D-glucosyl-beta-sitosterols were noticed (2001)
Anti-inflammatory A reduction in granuloma weight of 71.66% in the chronic Patil and Patil
study and 75.90% in acute inflammation confirmed the (2011)
anti-inflammatory properties of fig oil
Anti-diabetic A hypoglycemic effect was observed in rats on treatment Perez et al.
with fig oil (1998)
Immunomodulatory Amelioration of both humoral and cellular antibody Patil et al.
responses was observed (2010)
Sperm parameters An improvement in gonadosomatic index, sperm count, Naghdi et al.
and non-progressive spermatozoa motility was found with (2016)
fig oil ingestion
Adapted from (Rahmani and Aldebasi (2017)
Table 16.6 Volatile Organic Compounds identified through GC/MS in fig oils from three ripening stages and cultivars of figs
364

Taamriwthe Azegzaw Averkane

Odor threshold in Fully Fully Fully
Volatile Compounds Aroma characteristics water (μg/kg) Unripe Ripe ripe Unripe Ripe ripe Unripe Ripe ripe
Alcohol
Pent-1-en-3-ol Pungent link green vegetable and 400 nd 0.034 nd 0.035 0.026 0.020 nd nd 0.035
fruity nuances
Hexan-1-ol Pungent, ethereal, fruity, and 500 nd nd nd 0.026 0.024 0.024 nd nd nd
alcoholic
(E) hex-2-en-1-ol Fruity and herbal nuances 100 nd nd nd nd 0.020 nd nd nd nd
Benzyl alcohol Sweet, floral, fruity, and chemical 10,000 nd nd nd 0.166 nd nd nd nd nd
nuances
Pentan-1-ol Pungent, fermented bready, yeasty, nd nd nd nd nd nd nd nd 0.020 nd
moldy
3-methylbutan-1-ol Pungent, ethereal, fruity, cognac nd nd 0.050 0.085 0.028 nd 0.037 nd nd 0.032
and alcoholic
2-ethylhexan-1-ol Citrus, fresh, floral, oily and sweet nd 0.020 nd nd 0.050 0.054 0.028 nd nd 0.033
Aldehydes
Acetaldehyde Pungent, ethereal, fresh, fruity 15 0.124 0.148 0.273 0.075 0.064 0.126 nd 0.058 0.164
Heptanal Fresh, citrus, green odor 3 nd nd 0.032 nd nd nd nd nd nd
Nonanal Fresh, green lemon, citrus 1-98 nd 0.02 nd 0.026 nd 0.026 nd nd nd
Benzaldehyde Fruity, almond, nutty, powdery 350 nd 0.093 0.075 0.937 0.424 0.35 nd nd 0.09
Hexanal Fatty, green, vegetative, leafy, 5 0.311 0.553 0.359 0.788 0.33 0.258 0.226 0.152 0.141
fruity and woody nuance
(E)-2-methylbut-2- Strong green fruit nd 0.102 0.157 0.078 nd nd nd nd nd nd
enal
(E)-hex-2-enal Aldehydic fatty cheese, green 17-316 0.115 0.214 0.311 nd nd nd nd 0.05 0.175
banana
(E)-oct-2-enal Fatty green herbal banana, fresh 3 nd nd nd 0.016 nd nd nd nd nd
E. G. M. G. Rajendran

cucumber, waxy green leaf

Esters
Ethyl acetate Ethereal, sweet, fruity, grape and 5000 0.216 0.159 0.135 0.137 nd nd nd 0.198 0.333
rum-like
Methyl hexanoate Fermented, fusel, banana, fruity, 84 nd 0.093 nd nd 0.035 nd nd nd nd
ethereal
Methyl octanoate Sweet, green, vegetative, orange nd nd nd nd nd 0.026 nd nd nd 0.067
nuances
Ethyl Berry, fruity, fresh tropical nuances nd nd nd 0.037 nd nd nd nd nd nd
2-methylbutanoate
16 Fig (Ficus carica) Seed Oil

Ethyl hexanoate Fruity, sweet, pineapple 14 nd nd 0.034 nd nd nd nd nd nd

Terpenes
Limonene Citrus orange fresh, sweet 10 nd nd nd nd 0.215 0.109 nd nd nd
Linalool Floral, citrus, orange 6 nd 0.021 nd 1.049 0.498 0.892 nd nd 0.043
Epoxylinalol Floral honey nd nd 0.025 nd nd nd 0.028 nd nd nd
α-Santalene Woody nd nd 0.033 nd nd nd nd nd nd nd
1,8-cineol Herbal eucalyptus, camphor 1,3-12 nd nd nd nd nd nd 0.03 nd nd
medicinal
β-caryophyllene Sweet, spice, woody, clove dry 150 0.292 0.254 nd nd nd nd nd nd nd
Ketones
Acetone Ethereal, solvent, apple, pear nd nd 0.059 0.1 0.034 0.05 0.045 0.116 0.079 0.043
Heptan-2-one Fruity, cheese, ketonic, creamy 140 nd nd 0.073 nd nd nd 0.107 nd 0.152
nuances
6-methylhept-5-en-2- Apple, fruity, ketonic, musty, 160 nd nd nd 0.022 0.022 nd nd nd nd
one cheesy nuances
nd not detected
Description for Aroma was obtained from (https://www.thegoodscentscompany.com/); thresholds were obtained from Tamura et al. (2001); Yang et al. (2019);
Noguerol-Pato et al. (2012); Buttery et al. (1969); Wu et al. (2016); Takeoka et al. (1990); Czerny et al. (2008). Volatile compounds are classified according to
chemical classes. The mean ± SD of three samples in triplicate were analyzed and obtained from, Zidi et al. (2021).
365
366 E. G. M. G. Rajendran

classified according to visual maturity characteristics based on fruit firmness and

color change from Taamriwthe (TH), Azegzaw (AZ), and Averkane (AV) cultivars.
A significant increase in aldehyde content was observed in TH and AV fig oils.
AZ had a decline in them. Totally six aldehydes were observed in these fig oils. The
extract of terpenes was found only at the ripe stages in the TH cultivar. Linalool was
observed to be the most abundant terpene in the AZ cultivar. AV fig oils contained
only two terpenes. Esters were significantly increased obtaining from figs during the
ripening process for oil extraction. TH and AV had increased total ester, whereas AZ
had overall less proportion of esters. A significant increase was observed in alcohol
content during the ripening process of AV cultivars. Whereas AZ had a significant
decrease in alcohol content, and TH had only three alcohol compounds that showed
a significant increase in the oils obtained from fully ripe figs. A total of three ketones
were observed in this study, and only two were found in all three cultivars.

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Chapter 17
Composition and Functional Properties
of Fig (Ficus carica) Phenolics

Mustafa Kiralan, Onur Ketenoglu, Sündüz Sezer Kiralan,

and Fatih Mehmet Yilmaz

1 Introduction

Fig (Ficus carica L.) is a member of the Moraceae family and grown in warm, dry
climates (Lo Turco et al., 2020). According to the official data belonging to 2019,
the total world fig production is 1,315,588 tonnes. Turkey (310,000 tonnes), Egypt
(225,295 tonnes), Algeria (114,092 tonnes), Iran (130,328 tonnes), and Morocco
(153,472 tonnes) are the major producer countries, yielding almost 80% of the total
production (FAO, 2021). Ficus carica Linn is the best-known member of the Ficus
genus, known by over 135 names. It has been the subject of worldwide studies due
to the positive effects of its various biological activities on the endocrine system,
respiratory system, gastrointestinal system, and infectious diseases (Badgujar et al.,
2014). Figs are processed into wine, liquor, fruit juice, and jelly products and are
used fresh and dried (Backes et al., 2020).
Fig is an important constituent of a Mediterranean diet due to its nutritional value
and phytochemicals. Fig is an essential source of dietary fiber, amino acids, carbo-
hydrates, minerals, organic acids, and phenolic compounds (Slatnar et al., 2011;
Slavin, 2006; Wojdyło et al., 2016). Additionally, fig fruits are important sources of
vitamins (Vitamin C, thiamin, riboflavin) and trace minerals (potassium, calcium,

M. Kiralan (*) · S. S. Kiralan

Department of Food Engineering, Faculty of Engineering, Balikesir University,
Balikesir, Turkey
O. Ketenoglu
Department of Food Engineering, Faculty of Agriculture, Eskisehir Osmangazi University,
Eskisehir, Turkey
F. M. Yilmaz
Department of Food Engineering, Faculty of Engineering, Aydin Adnan Menderes University,
Aydin, Turkey

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 369
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_17
370 M. Kiralan et al.

magnesium). Furthermore, figs are cholesterol-free and contain many essential

amino acids, fiber, and antioxidant compounds (Görgüç et al., 2021). Antioxidant
compounds such as phenolics (phenolic acids, flavonoids, carotenoids) are the
major constituents that provide the fig fruits‘characteristic color, flavor, and aroma
(Fateh Aljane & Ferchichi, 2009; Arvaniti et al., 2019). The overall composition of
fresh fig is summarized in Table 17.1. Considering dried figs, total carbohydrate, fat,
protein, dietary fiber, and moisture content were 73.5%, 0.56%, 4.67%, 3.68%, and
16.6%, respectively (Soni et al., 2014). Therefore, dried and fresh figs are good
carbohydrates and dietary fiber sources.
Besides macronutrients, fig contains different phytochemicals such as anthocya-
nins, polyphenols, and carotenoids (Jagtap & Bapat, 2020). Phenolic compounds
are secondary metabolites synthesized through the shikimic acid and phenylpro-
panoid pathways (de la Rosa et al., 2019). Fruits are rich in phenolic compounds
that contribute to the prevention of several degenerative diseases (Renard, 2018).
The health effects of phenolic compounds are largely attributed to the antioxidant
activity and other mechanisms of action (Shahidi et al., 2019). These activities are
closely related to the chemical diversity of phenolic compounds, especially the
hydroxyl groups on the aromatic rings. In addition, the structure of hydroxyl groups
attached to phenolic compounds influences their activities (Vuolo et al., 2019).
Figs exhibit numerous biological activities such as antioxidant, antimicrobial,
and anti-diabetic (Arvaniti et al., 2019; Benmaghnia et al., 2019; Oliveira et al.,
2009; Wojdyło et al., 2016), in particular, the different parts are reported to show
antioxidant (Ahmad et al., 2013; Mostapha Bachir Bey & Louaileche, 2015;
Solomon et al., 2006), antimicrobial (Aref et al., 2010; Jeong et al., 2009; Lazreg-
Aref et al., 2012; Mahmoudi et al., 2016), anticancer (Jasmine et al., 2015;
Purnamasari et al., 2019; Rubnov et al., 2001), and antidiabetic (Zaffer Ahmad
et al., 2013) activities. The phenolic compounds, which are mainly responsible for
these biological activities, have recently gained considerable attention, and the phe-
nolic content of fig changes depending on various factors such as fruit quality, pro-
cessing, and extraction techniques. This chapter discusses the composition of
phenolic compounds and the biological activities of fig phenolics.

Table 17.1 The composition of some nutritients in fresh fig (g/100 g)

Nutrition Quantity
Water 79.1
Protein 0.75
Total lipid (fat) 0.3
Carbohydrate 19.2
Fiber, total dietary 2.9
Sugars 16.3
Source: U.S. Department of Agriculture; FDC ID: 173021; NDB Number: 9089; FDC:
Published:4/1/2019
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 371

2 Total Phenolics, Total Flavonoids and Total

Anthocyanins in Figs

Fig fruits contain phenolic acids such as chlorogenic acid, flavanols such as (+)-
catechin and (−)-epicatechin, flavonols such as quercetin-3-O-glucoside,
kaempferol-3-O-glucoside, and rutin, flavons such as luteolin-8-C-glucoside, and
anthocyanins such as cyanidin-3-O rutinoside (Slatnar et al., 2011). A general rep-
resentation of fig phenolics is given in Fig. 17.1.
Total phenolics are determined using a colorimetric method called Folin-
Ciocalteu (F-C) assay. This method is based on electron transfer reactions between
the F-C reagent and polyphenols (Ainsworth & Gillespie, 2007; Sánchez-Rangel
et al., 2013).
A summarizing list of total phenols in fresh and dried figs is given in Table 17.2,
and total flavonoids and total anthocyanins in figs, depending on the part, and the
origin of the fruit, are summarized in Table 17.3. The part of the fruit is one of these
factors affecting the phenolic content. In general, leaves have higher total phenols
values than other parts such as peels and pulps (Oliveira et al., 2009). These differ-
ences are attributed to especially flavonoids which have photoprotection properties
in plants (Treutter, 2006). Besides, the skin part of figs contains a higher content of
total phenols than the extracts from the pulp part. Solomon et al. (2006) explain
these differences and emphasize that anthocyanins are the most abundant phenolics
in the skin part of figs, so they contribute to the total phenolics and phytochemicals.

Fig. 17.1 The main phenolic compounds determined in fig fruits

372 M. Kiralan et al.

Table 17.2 Total phenols in fresh and dried figs

Sample
(Part/
Tissue) Variety, origin, color Assay method Concentration References
Fruit 76 local Mediterranean Spectrometric- 28.6–211.9 mg Çalişkan &
(whole) fig accessions from Folin- GAE/100 g FW Aytekin Polat
Turkey Ciocalteau (2011)
Leaves Spectrometric- 4.363–6.909 mg/ dry Konyalιoğlu
Folin- mass et al. (2005)
Ciocalteau
Peel White varieties of F. HPLC-DAD 600.6, 723 mg/kg Oliveira et al.
Pulp carica (Pingo de Mel lyophilized extract (2009)
Leaves and Branca (Peel)
Tradicional) from 36.6, 130.3 mg/kg
Portugal lyophilized extract
(pulp)
32412.0, 34473.6 mg/
kg lyophilized extract
(leaves)
Fruit Škofjotka (with the HPLC-DAD Miljska cultivar had Veberic et al.
(whole) synonym higher values in total (2008)
‘Zuccherina’) a white phenol content, while
type fruit, ‘Črna Škofjotka cultivar is
petrovka’ and ‘Miljska poor
figa’, both dark type
fruit from Slovenia
Skin and Eighteen fig cultivars HPLC-DAD 19.1–140.2 mg/ Vallejo et al.
pulp from Spain mg/100 g FW in fig (2012)
skins
0.0–11.3 mg/100 g FW
in fig pulps
Fruit, Mission, Chechick, Spectrometric- 48.6–281.1 mg of Solomon et al.
skin and Bursa (dark purple) Folin- GAE/100 g of FW in (2006)
pulp Brown Turkey Ciocalteau fruits
(purple), Brunswick 41.7–463 mg of
(red yellow), and GAE/100 g of FW in
Kadota (green) skins
36.5–100.6 mg of
GAE/100 g of FW in
pulps
(continued)
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 373

Table 17.2 (continued)

Sample
(Part/
Tissue) Variety, origin, color Assay method Concentration References
Leaves Ten Algerian cultivars Spectrometric- 42.889–58.704 mg Mahmoudi et al.
Onk Elhamam Folin- GAE/g DE (2016)
Hamra Ciocalteau
Zarrouk
Boughandjo
Safra
Bidha
Chatwi
Bither
Bakkor
Dhokkar
Whole, Two Turkish cultivars Spectrometric- Sarilop-yellow Kamiloglu &
skin, Sarilop-yellow Folin- Skin-351 mg GAE Capanoglu
pulp Bursa siyahi- purple Ciocalteau /100 g FW (2015)
Pulp-167 mg GAE
/100 g FW
Whole-211 mg GAE
/100 g FW
Bursa siyahi- purple
Skin-930 mg GAE
/100 g FW
Pulp-351 mg GAE
/100 g FW
Whole-493 mg GAE
/100 g FW
Whole 24 Turkish fig Spectrometric- 24–237 mg GAE Ercisli et al.
genotypes and two Folin- /100 g FW (2012)
standard fig (Ficus Ciocalteau
carica L.) cultivars
(Sarilop and Bursa
Siyahi)
Whole Ten different Spanish UPLC-PDA-FL 715 to 186 mg/100 g dm Wojdyło et al.
fig cultivars (Ficus (2016)
carica L.)
Whole 27 Tunisian ecotypes Spectrometric- 51.50–100.23 mg GAE F Aljane et al.
Folin- /100 g FW (2020)
Ciocalteau
Whole 11 local and Spectrometric- 90.75–524.74 mg GAE Hssaini et al.
introduced cultivars Folin- /100 g DW (2020)
from Morocco Ciocalteau
(continued)
374 M. Kiralan et al.

Table 17.2 (continued)

Sample
(Part/
Tissue) Variety, origin, color Assay method Concentration References
Dried Tounsi and Temri Spectrometric- Tounsi cultivar Viuda-Martos
Leaves cultivars from Tunisia Folin- Leaves-618.29 mg et al. (2015)
and fruits Ciocalteau GAE /100 g DW
(whole) Fruits-200.18 mg GAE
/100 g DW
Temri cultivar
Leaves-686.88 mg
GAE /100 g DW
Fruits-124.48 mg GAE
/100 g DW
Latex 18 diferent cultivars of Spectrometric- The maceration Shahinuzzaman
F. carica from Folin- extraction method et al. (2020)
Malaysia Ciocalteau 233.14–311.83 μg
GAE/mL
Ultrasound-assisted
Extraction
(224.37–291.98 μg
GAE/mL
FW Fresh weight, GAE gallic acid equivalents, DE dry plant extract, dm dry matter

Additionally, location is another influential factor in total phenols. According to

Kamiloglu and Capanoglu (2015), the figs that grew in Turkey generally had higher
total phenolic values than in other countries.
Latex is another important part of the fig, and it contains various phenolic com-
pounds. Shahinuzzaman et al. (2020) extracted phenolic compounds from 18 differ-
ent fig cultivars using two different extraction techniques: maceration and
ultrasound-assisted extraction (UAE) with 75% ethanol. The latex was reported as a
valuable natural source of phenolics. The latex from the White Genoa cultivar
included higher amounts of total phenols.
Regarding color, the extracts from dark-colored fig cultivars contain higher lev-
els of total phenols than the light-colored fig cultivars. For example, Solomon et al.
(2006) stated that a dark-colored variety of Mission contained a high content of total
phenols (281 mg/100 g of fresh weight (FW)), while a green variety, namely Kadota,
was poor for total phenols (49 mg/100 g of FW).
Ripening is another factor in the phenolics of figs. Pereira et al. (2017) found that
concentrations of phenolics increased during the ripening process. However, differ-
ences in the concentrations of total phenols were observed during ripening.
Moreover, the effect of ripening on total phenolics of dark- and brown-colored cul-
tivars was stronger than green- and yellow-green-colored cultivars. This effect could
be related to the changes of anthocyanins accumulated in the skin during the ripen-
ing stage of dark-colored cultivars.
The drying process is also another essential factor in the total phenols of figs. As
seen in Table 17.4, the content of total phenolics, total flavonoids, and total
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 375

Table 17.3 Total flavonoids and total anthocyanins in figs

Sample
(Part/
Tissue) Variety, origin, color Total flavonoids Total anthocyanins References
Leaves Ten Algerian cultivars 11.667–16.211 mg – Mahmoudi
Onk Elhamam QE/g DE et al. (2016)
Hamra
Zarrouk
Boughandjo
Safra
Bidha
Chatwi
Bither
Bakkor
Dhokkar
Whole, Two Turkish cultivars Sarilop-yellow Sarilop-yellow Kamiloglu &
skin, pulp Sarilop-yellow 7–63 mg of CE/100 g 0.8–5.9 mg of Capanoglu
Bursa siyahi- purple DW C3G/100 g DW (2015)
Bursa siyahi- purple Bursa
66–234 mg of siyahi- purple
CE/100 g DW /100 g DW
Whole 76 local – 0.0–298.9 μg C3R Çalişkan et al.
Mediterranean fig /g FW (2011)
accessions from
Turkey
Whole 24 Turkish fig – 0–42 μg C3R /g Ercisli et al.
genotypes and two FW (2012)
standard fig (Ficus
carica L.) cultivars
(Sarilop and Bursa
Siyahi)
Whole Ten different Spanish – 0.4– Wojdyło et al.
fig cultivars (Ficus 122 mg/100 g dm (2016)
carica L.)
Whole 27 Tunisian ecotypes 0.33–17.59 mg 1.61–11.67 mg Aljane et al.
QE/100 g FW C3G/ 100 g FW (2020)
Whole 11 local and 14.59– 3.1–24.8 mg Hssaini et al.
introduced cultivars 83.71 mg CE/100 g C3R/100 g DW (2020)
from Morocco DW
QE quercetin equivalents, DE dry plant extract, FW fresh weight, C3G cyanidin-3-glycoside, C3R
cyanidin -3-rutinoside, QE quercetin, CE Catechin

anthocyanins differ in the literature. Slatnar et al. (2011) used two different drying
processes: sun-drying and oven-drying, to obtain dried figs. The total phenolic com-
pounds were 74.9 mg GAE/kg FW in fresh figs to 530 mg GAE/kg FW in dried figs.
The higher values for total phenols were observed in the samples using the oven-
drying process compared to sun-drying. Kamiloglu and Capanoglu (2015) found a
decrease in total phenols of yellow and purple figs after the sun-drying process (8
and 15%, respectively). Their study evaluated total flavonoids, total
Table 17.4 Effect of drying on the content of total phenols, total flavonoids and total anthocyanins of figs
Total phenols Total flavonoid content Total anthocyanin content References
Dried Dried Dried
Sample Variety,
(part/ origin, Oven- Oven- Oven-
tissue) color Fresh Sun-drying drying Lyophilized Fresh Sun-drying drying Lyophilized Fresh Sun-drying drying
Whole Sarilop- 211 mg of 193 mg of – – 8 mg of CE/100 g 14 mg of – – 4.6 mg of 0.1 mg of – Kamiloglu
yellow GAE/100 g DW) GAE/100 g DW CE/100 g C3G/100 g C3G/100 g and
DW) DW DW DW Capanoglu
(2015)
Whole Bursa 493 mg of 417 mg of – – 66 mg of 52 mg of – – 83 mg of 14.5 mg of – Kamiloglu
siyahi- GAE/100 g DW) GAE/100 g CE/100 g DW CE/100 g C3G/100 g C3G/100 g and
purple DW) DW DW DW Capanoglu
(2015)
Whole 198.81–307.64 mg 81.77– – – 67.33–147.51 mg 32.78– – – – – – Nakilcioğlu
GAE/100 g DM 212.36 mg RE/100 g DM 52.52 mg and Hışıl
GAE/100 g RE/100 g (2013)
DM DM
Whole Aberkane 107.08– – – 30.81– 84.63– – – 12.09– – – – Mostapha
(black), 181.68 mg/100 g 41.91 mg/100 g 151.47 mg/100 g 20.82 mg/100 g Bachir Bey
Azandjar, DM DM DM DM et al. (2017)
and
Bouankik
(dark-
purple)
DW Dry weight, GAE gallic acid equivalents, CE (+)-catechin equivalent, C3G cyanidin-3-glycoside
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 377

proanthocyanidin content, and total anthocyanin content depending on the sun-

drying process. There were no significant changes between the total flavonoid con-
tent of fresh and dried fruits of purple figs. However, dried fruits from yellow figs
contained more flavonoids than the fresh ones. The drying process showed a
decrease in total proanthocyanidins in purple figs, while an increase was determined
for yellow figs. The highest decrease was observed in the total anthocyanin content
for purple and yellow figs depending on the sun-drying process (83% in purple figs,
98% in yellow figs). Lachtar et al. (2021) used two different systems: direct solar
drying (open-air) and solar drying in a greenhouse to dry Bither Abiadh and Bidhi
fig cultivars from Tunisia. The two cultivars dried in a greenhouse had higher total
phenols than open air-dried samples. Mostapha Bachir Bey, Richard, Meziant,
Fauconnier, and Louaileche (Bey et al., 2017) found a drastic reduction in total
phenols and total flavonoids of figs dried by lyophilization.
Besides drying, the freezing process negatively influences the total phenol, total
flavonoid, and total anthocyanins. A dramatic decrease was observed in the total
phenol, total flavonoid, and total anthocyanins of the frozen figs in comparison to
fresh figs, where the concentration of these phytochemicals in fresh figs was 31.0 mg
GAE/100 g, 11.4 mg QE/100 g, and 0.8 mg C3G/100 g, while in frozen figs their
concentrations were the lowest (14.0 mg GAE/100 g, 3.6 mg QE/100 g, and not
detected, respectively) (Petkova et al., 2019).

3 Individual Phenolic Compounds in Figs

Individual phenolic compounds exhibit a wide variety in figs, and their amount
changes with different fruit cultivars, color, ripening, and drying methods. Table 17.5
summarizes the most common individual phenolics in fresh and dried samples.
Wojdyło et al. (2016) identified 11 polyphenolic compounds in such chemical
classes as flavan-3-ols, phenolic acids, flavonols, flavons, and anthocyanins from
different fig cultivars grown under the same conditions in Spain. The predominant
flavanol was quercetin-3-O-rutinoside, with the highest concentration of
328 mg/100 g dry matter (DM) in the Verdal brevas cultivar. Chlorogenic acid was
the only phenolic acid present in fig fruits, and the highest concentration of this
compound was reported as 124 mg/100 g DM in Verdal fig. Catechin was a major
compound among flavon-3-ols with a 20.6 to 95.7 mg/100 g DM concentration.
Cyanidin-3-O-rutinoside was the main anthocyanin in figs, and Cuello Dama Negra
cultivar contained the highest amount (116 mg/100 g DM).
Kamiloglu and Capanoglu (2015) studied individual phenolics of fruits with dif-
ferent colors. The predominant flavanoid was rutin for yellow figs. (12 mg/100 g
DM), whereas cyanidin-3-rutinoside was found as the major anthocyanin for purple
figs. (36 mg/100 g DM).
Faleh et al. (2012) evaluated the phenolic composition of three Tunisian fig phe-
notypes with different colors: black, green, and red. Quercetin-3-O-rutinoside was
the major phenolic in all phenotypes, and its concentration was higher in the red
378 M. Kiralan et al.

Table 17.5 The most common individual phenolics in fresh and dried samples
Subclass Component Dried Fresh References
Phenolic Chlorogenic acid 1.4–2.0 mg/100 g 2–5.8 mg/100 g Vallejo et al.
acids FW (skin) (2012)
nd-1.3 mg/100 g
FW (pulp)
0.46– Veberic et al.
1.71 mg/100 g FW (2008)
(fruit)
3.42– 1.33– Slatnar et al.
15.88 mg/100 g 4.91 mg/100 g FW (2011)
(sun-drying)
13.96–
32.42 mg/100 g
(oven-drying)
8.8– – Wojdyło et al.
124.5 mg/100 g (2016)
DM
0.40– 2.97– Nakilcioğlu and
2.21 mg/100 g DM 6.54 mg/100 g Hışıl (2013)
DM
0.8–1.1 mg/100 g 1.9–8 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
12–21 mg/100 g
DM (skin)
0.5–6 mg/100 g
DM (pulp)
0.06– – Pourghayoumi
0.55 mg/100 g DM et al. (2016)
Gallic acid 0.14– Veberic et al.
0.38 mg/100 g FW (2008)
(fruit)
0.47– 1.15– Nakilcioğlu and
2.18 mg/100 g DM 6.98 mg/100 g Hışıl (2013)
DM
0.1–0.5 mg/100 g 0.3–0.3 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
0.2–0.3 mg/100 g
DM (skin)
0.3–0.5 mg/100 g
DM (pulp)
Syringic acid 0.022– Veberic et al.
0.104 mg/100 g (2008)
FW (fruit)
0.41– 3.03– Nakilcioğlu and
2.83 mg/100 g DM 4.45 mg/100 g Hışıl (2013)
DM
(continued)
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 379

Table 17.5 (continued)

Subclass Component Dried Fresh References
Ellagic acid 0.2–0.3 mg/100 g 0.2–0.2 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
0.3–0.7 mg/100 g
DM (skin)
0.1–0.2 mg/100 g
DM (pulp)
p-Coumaric acid 1.1–3.1 mg/100 g 2.7–4.7 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
5.1–18 mg/100 g
DM (skin)
2.2–3.2 mg/100 g
DM (pulp)
Flavones Luteolin – 0.1–1.9 mg/100 g Vallejo et al.
6C-hexose-8C- FW (skin) (2012)
pentose
Apigenin- – 0.8–2.4 mg/100 g Vallejo et al.
rutinoside FW (skin) (2012)
Apigenin-C- 0.5–24.3 mg/100 g – Wojdyło et al.
hexoside-pentoside DM (2016)
Flavonols Kaempferol- 1.0–2.0 mg/100 g 0.2–0.9 mg/100 g Vallejo et al.
rutinoside FW (skin) (2012)
0.2–0.9 mg/100 g 0.3–0.3 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
1.4–4 mg/100 g
DM (skin)
nd-0.1 mg/100 g
DM (pulp)
Quercetine- 2.1–2.8 mg/100 g Nd-1.7 mg/100 g Vallejo et al.
acetilglucoside FW (skin) (2012)
Quercetin- 10.2– 2.9–15.8 mg/100 g Vallejo et al.
rutinoside 13.0 mg/100 g FW (skin) (2012)
Nd-1.8 mg/100 g
FW (pulp)
Quercetin- 0.2–3.2 mg/100 g Vallejo et al.
glucoside FW (skin) (2012)
Rutin – 4.89– Veberic et al.
28.7 mg/100 g FW (2008)
(fruit)
1.38– 0.61– Slatnar et al.
12.06 mg/100 g 1.86 mg/100 g FW (2011)
(sun-drying)
3.75–
14.62 mg/100 g
(sun-drying)
5.75– 15.16– Nakilcioğlu and
12.35 mg/100 g 68.21 mg/100 g Hışıl (2013)
DM DM
(continued)
380 M. Kiralan et al.

Table 17.5 (continued)

Subclass Component Dried Fresh References
9–15 mg/100 g 12–13 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
86–180 mg/100 g
DM (skin)
0.9–2 mg/100 g
DM (pulp)
Kaempferol-3-O- 0.31– 0.04– Slatnar et al.
glucoside 0.59 mg/100 g 0.13 mg/100 g FW (2011)
(sun-drying)
0.56–
1.43 mg/100 g
(oven-drying)
Luteolin-8-C- 0.13– – Slatnar et al.
glucoside 0.16 mg/100 g (2011)
(sun-drying)
0.21–
0.45 mg/100 g
(oven-drying)
Quercetin-3-O- 0.56– 0.18– Slatnar et al.
glucoside 3.55 mg/100 g 0.60 mg/100 g FW (2011)
(sun-drying)
1.10–
2.98 mg/100 g
(oven-drying)
0.2–0.7 mg/100 g 0.3–1.2 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
1.9–9.4 mg/100 g
DM (skin)
0.1–0.2 mg/100 g
DM (pulp)
0.12– Pourghayoumi
1.27 mg/100 g DM et al. (2016)
Keampferol-3-O- 0.6–30.5 mg/100 g – Wojdyło et al.
rutinoside DM (2016)
Quercetin-3-O- 39.0– – Wojdyło et al.
rutinoside 328.3 mg/100 g (2016)
DM
4.33–88.40 mg/kg Faleh et al.
DM (2012)
Quercetin-3- 4.6–14.2 mg/100 g – Wojdyło et al.
Galactoside DM (2016)
Quercetin-3-O- 5.3–58.0 mg/100 g – Wojdyło et al.
malonyl- DM (2016)
galactoside
Anthocyanins Cyanidin-3- Nd-11.1 mg/100 g Vallejo et al.
glucoside FW (skin) (2012)
(continued)
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 381

Table 17.5 (continued)

Subclass Component Dried Fresh References
Nd-0.1 mg/100 g 0.1–6 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
Nd-23 mg/100 g
DM (skin)
0.1–2.8 mg/100 g
DM (pulp)
Cyanidin-3- Nd- Vallejo et al.
rutinoside 108.9 mg/100 g (2012)
FW (skin)
Nd-9.5 mg/100 g
FW (pulp)
0.12– 0.21– Slatnar et al.
0.26 mg/100 g 0.62 mg/100 g FW (2011)
(sun-drying)
0.12–
0.31 mg/100 g
(oven-drying)
Nd-1.5 mg/100 g 2.1–36 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
0.1–180 mg/100 g
DM (skin)
2.2–18 mg/100 g
DM (pulp)
Cyanidin-3,5-O- 0.0–1.9 mg/100 g – Wojdyło et al.
diglucoside DM (2016)
Pelargonidin-3-O- 0.0–3.8 mg/100 g – Wojdyło et al.
rutinoside DM (2016)
Cyanidin-3-O- 0.4– Wojdyło et al.
rutinoside 116.2 mg/100 g (2016)
DM
Flavanols (−)-Epicatechin 0.34– Veberic et al.
0.97 mg/100 g FW (2008)
(fruit)
10.44– 7.11– Slatnar et al.
23.30 mg/100 g 8.67 mg/100 g FW (2011)
(sun-drying)
26.66–
36.65 mg/100 g
(oven-drying)
9.0–43.1 mg/100 g Wojdyło et al.
DM (2016)
14.76– 42.29– Nakilcioğlu and
23.42 mg/100 g 76.90 mg/100 g Hışıl (2013)
DM DM
(continued)
382 M. Kiralan et al.

Table 17.5 (continued)

Subclass Component Dried Fresh References
0.6–2.2 mg/100 g 1.1–6.9 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
2.7–29 mg/100 g
DM (skin)
0.9–4.8 mg/100 g
DM (pulp)
(+)-Catechin – 1.07– Veberic et al.
4.03 mg/100 g FW (2008)
(fruit)
5.88– 1.36– Slatnar et al.
11.46 mg/100 g 2.88 mg/100 g FW (2011)
(sun-drying)
15.57–
19.75 mg/100 g
(oven-drying)
20.6– – Wojdyło et al.
95.7 mg/100 g DM (2016)
1.5–2.5 mg/100 g 2.3–13 mg/100 g Kamiloglu and
DM DM (whole) Capanoglu (2015)
6–8 mg/100 g DM
(skin)
2.1–27 mg/100 g
DM (pulp)
1.25– Pourghayoumi
4.77 mg/100 g DM et al. (2016)

phenotypes (12.3–88.4 mg/kg DM) than in the green (4.33–55.5 mg/kg DM) and
black ones (17.5–39.6 mg/kg DM).
Vallejo et al. (2012) evaluated the individual phenolics from different parts of fig.
According to the results, cyanidin-3-rutinoside was found as a major phenolic com-
pound among anthocyanins in the skin part of fig, reaching up to 108.9 mg/100 g,
while fig pulp was poor in cyanidin-3-rutinoside (9.5 mg/100 g).
Kamiloglu and Capanoglu (2015) evaluated the individual phenolics for skin and
pulp parts of figs. The skin part was rich in chlorogenic acid, ellagic acid, and p-
coumaric acid than pulp, whereas gallic acid was found at a higher pulp concentra-
tion than skin. Rutin was the predominant flavonol, identified mainly in the fruit
skins. Among flavon-3-ols, catechin was found mainly in the skin part of yellow figs
and the pulp of purple figs. The skin part contained higher anthocyanin levels than
the purple figs pulp, with eight and ten folds of cyanidin-3-glucoside and cyanidin-3-
rutinoside, respectively.
The ripening stage is another factor that affects the individual content of pheno-
lics in figs. Slatnar et al. (2011) evaluated the effect of sampling date (ripening
stage) on individual phenolics. The highest content of individual phenolics was
observed at the second sampling of the first crop. Epicatechin was the predominant
individual phenolic compound, with a concentration ranging between 7.11 and
8.67 mg 100 g FW in fresh figs.
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 383

The rain-fed condition also affects the phenolic composition of figs. Sedaghat
and Rahemi (2018) found that (+) catechin, chlorogenic acid, (−) epicatechin, and
quercetin-3-O- glucoside exhibited higher values in rain-fed figs during fruit ripen-
ing. In the last ripening stage, the concentration of (+) catechin, Chlorogenic acid,
(−) epicatechin, and Quercetin-3-O-glucoside reached up to 17.06, 15.51, 33.3, and
4.19 mg/g, respectively.
Drying methods are also influential factors in the concentration of individual
phenolic compounds. Slatnar et al. (2011) reported that the oven-drying process
increased individual phenolics other than cyanidin-3-O-rutinoside compared with
the sun-drying process. Kamiloglu and Capanoglu (2015) reported that the chloro-
genic acid and p-coumaric acid contents decreased, while gallic acid increased after
drying. After drying, the decreasing trend was observed in all flavonols of yellow
figs. (24% for rutin, 33% both for kaempferol-rutinoside and Q3G, and 50% for
quercetin derivatives). A similar loss was observed in Q3G and quercetin derivatives
1 and 2 for purple figs. On the other hand, for purple figs, rutin and kaempferol
rutinoside increased with drying process (16% and 200%, respectively).

4 Extraction Methods

Several extraction methods, including conventional and novel techniques, have

been used to extract the phenolic compounds from the fig fruit. Maghsoudlou et al.
(2016) investigated the effects of different extraction techniques on the phenolic
compounds of fig. The authors evaluated the subcritical water extraction, ultrasound-
assisted extraction (UAE), and shaker solvent extraction techniques, and the anti-
oxidant activities of the extracted compounds were determined using DPPH·,
reducing power (RP), and rancimat tests. According to their results, subcritical
water extraction was the most appropriate technique for higher extraction abilities
for the total phenolics and flavonoids and obtaining the lowest values in the DPPH·
and RP tests (in the skin part). The authors also stated that the ultrasound-assisted
extraction gave better results when the Rancimat test was used to determine the
antioxidant activity of the extracts.
Alcántara et al. (2020) studied the effects of UAE and solvent on the phenolic
profile of fig leaves extracts. The researchers deployed the liquid solvent medium in
two ways: a) distilled water and b) 50% (v/v) of water-alcohol solution. In compar-
ing UAE and conventional extraction techniques, the researchers stated that UAE
extracted more carotenoids while the extraction performance of total phenolics
using conventional extraction was higher than the UAE. In conclusion, the authors
stated that the highest anti-inflammatory effect was observed when the extractions
were carried out using the hydro-ethanolic solvent.
Qin et al. (2015) used ionic liquid-based UAE to extract such phenolic compo-
nents as rutin, psoralen, chlorogenic acid, gallic acid, and bergapten from different
parts of fig. In addition, the researchers performed an optimization in their UAE
procedure. In conclusion, they stated that the highest extraction yields of the
384 M. Kiralan et al.

phenolic compounds were obtained at the optimal conditions of 1.0 M of 1-butyl-3-

methylimidazolium hexafluorophosphate [BMIM][PF6] concentration, the solid-
liquid ratio of 1:50 and the extraction temperature of 30 °C. Furthermore, according
to their findings, the ionic liquid usage achieved an increase in the extraction yield
and a decrease in the extraction time.
Jokić et al. (2014) also compared the use of UAE and classic solid-liquid extrac-
tion and determined their effects on the total phenolics, total flavonoids, and total
color change of fig extracts. Their results showed that higher total phenolic and
flavonoid concentrations (up to 13.7 and 18.5%, respectively) could be obtained
using the UAE. Besides, UAE achieved a decrease in the total color changes of the
fig extracts compared to classical extraction. The researchers concluded that higher
quality of extracts and higher extraction yields could be achieved by using UAE.

5 Biological Activities of Figs

5.1 Antioxidant Activity

The antioxidant effect of phenolic compounds is based on their capacity to reduce

oxygen concentration, which has a degenerative effect on the body, to prevent the
initiation of oxidation, and to scavenge free radicals (Karabulut & Yemiş, 2019;
Minatel et al., 2017). Several studies indicated that phenolic compounds found in
plant materials positively affect human health, attributed mainly to their antioxidant
activities (Bucić-Kojić et al., 2011). Dried figs are good sources of non-enzymatic
antioxidants that positively impact human health (Mostapha Bachir Bey &
Louaileche, 2015), and dark-colored fig varieties contain more polyphenols flavo-
noids, flavonols, anthocyanins, and proanthocyanidins than lighter varieties
(Mostapha Bachir Bey & Louaileche, 2015). The different antioxidant activities of
various extracts from figs depending on the cultivar and applied antioxidant tests are
presented in Table 17.6.
Slatnar et al. (2011) evaluated the antioxidant activity with the DPPH method in
different drying processes. Antioxidant activities of dried figs using the oven and
sun-drying methods exerted almost two-fold higher values than fresh figs.
F Aljane et al. (2020) examined the antioxidant activities of 27 Tunisian ecotypes
using DPPH· and ABTS assays. The inhibition percentage in the DPPH assay
changed between 11.3% for Besbessi and 64.7% for Bouharrag. The scavenging
activity of ABTS radical ranged from 263 mg equivalent vitamin C/100 g FW
(Mahdoui) to 676 mg equivalent vitamin C/100 g FW (Nemri).
Nakilcioğlu and Hışıl (2013) determined the antioxidant activities of ten fresh
figs from the Sarilop cultivar and also evaluated the effect of the drying process on
the antioxidant activity. The antioxidant activities of dried samples were higher than
fresh samples. DPPH· scavenging activity of fresh samples was between
1657–2241 mg Trolox equivalent/100 g DM, while it was found between
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 385

Table 17.6 Studies on the antioxidant activities of figs

Antioxidant test Fresh Fruit Dried Fruit References
ABTS – IC50 value: 19.8 mg/ Soni et al. (2014)
mL
263.7–676.13 mg – F Aljane et al.
equivalent vitamin (2020)
C/100 g FW
FRAP – 60.48 μg BHT Soni et al. (2014)
equivalent/mg
sample
4.6–18.7 mmol Fe2+/ kg – Ercisli et al.
FW (2012)
DPPH EC50: 0.72 mg/mL Yang et al. (2009)
(WE)
EC50:0.61 mg/mL
(PS)
Peel – Harzallah et al.
EC50: 4.52–26.7 mg/mL (2016)
Pulp
EC50: 10.59–15.45 mg/
mL
Whole fruit
EC50: 7.04–14.6 mg/mL
– IC50 value: Elahe
0.45–3.45 mg/mL Maghsoudlou
et al. (2017)
Inhibition percentage: – F Aljane et al.
11.37–64.73% (2020)
– 28.33% -45.25% Bey and
Louaileche (2015)
Superoxide radical EC50:0.95 mg/mL Yang et al. (2009)
scavenging activity (PS)
EC50:2.81 mg/mL
(WE)
137.42–315.70 mg Bey & Louaileche
GAE/100 g (2015)
Hydroxyl radical At the Yang et al. (2009)
scavenging activity concentration of
4 mg/mL
−43.4% for PS
−31.0% for WE
Hydrogen peroxide 92.21–210.77 mg Bey and
scavenging activity GAE/100 g Louaileche (2015)
Phosphomolybdenum 621.72–1024.37 mg Bey and
antioxidant activity GAE/100 g Louaileche (2015)
(continued)
386 M. Kiralan et al.

Table 17.6 (continued)

Antioxidant test Fresh Fruit Dried Fruit References
Reducing power At the Yang et al., 2009)
concentration of
4 mg/mL
– OD value: 0.58
for WE
– OD value: 0.35
for PS
IC50 value: Maghsoudlou
0.90–4.62 mg/ml et al. (2017)
ORAC 0.46–1.33 mM Wojdyło et al.
Trolox/100 g DM (2016)
TEAC – Whole Solomon et al.
25.0–716.3 μmol of (2006)
TE/100 g
Skin
82.0–1987.0 μmol
of TE/100 g
Pulp
20.8–357.5 μmol of
TE/100 g
36–623 μg Trolox – Ercisli et al.
Equivalent/100 g (2012)
1.44–3.14 μmol TE/g – Crisosto et al.
FW (2010)
WE water extract, PS crude hot-water soluble polysaccharide

663.2–1103 mg Trolox equivalent /100 g DM in dried samples. FRAP scavenging

capacity of fresh samples was between 553.4–914.6 mg FeSO4/100 g DM, while the
scavenging capacity of dried samples changed between 176.49 and 290.94 mg
FeSO4/100 g DM.
Yang et al. (2009) used different antioxidant assays such as DPPH·, superoxide
radical scavenging activity, hydroxyl radical scavenging activity, and RP to evaluate
the activities of water extract (WE) and crude hot-water soluble polysaccharide (PS)
from fig fruit. Both WE and PS showed good scavenging activities on DPPH· (EC50
values for WE and PS were 0.72 and 0.61 mg/ml, respectively). Besides, PS had
higher antioxidant activity than WE on superoxide radical, hydroxyl anion radical,
and reducing power assays.
Harzallah et al. (2016) examined the antioxidant properties of juices of peel,
pulp, and total fruit of figs from three different varieties with three different colors
(green, purple, and black) grown in Tunisia via DPPH· scavenging capacity and
RP. Black peel juice showed the highest antioxidant activities (DPPH and RP), fol-
lowed by black fig total fruit juice.
Pereira et al. (2017) reported that the hydrophilic fraction had higher antioxidant
activity than the lipophilic fraction in the skin and flesh parts from nine different
cultivars. Considering the parts of the fruit, the skin part had higher values (between
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 387

two and ten times) than the flesh, depending on the cultivar. Besides, the dark-
colored cultivars showed higher antioxidant activity (16.3–177 mmol Trolox equiv-
alents per 100 g of FW) than the lighter ones. Solomon et al. (2006) emphasized that
the effect of the skin part (82–2000 μmol/100 g of FW) on antioxidant activity was
higher than that of the pulp (21–358 μmol/100 g of FW).
Besides fruit, latex is also rich in antioxidant compounds. Shahinuzzaman et al.
(2020) extracted bioactive compounds from fig fruit using maceration (ME) and
ultrasound-assisted extraction (UAE) methods. Amongst the 18 cultivars collected
in Malaysia, the latex from the White Genoa cultivar exhibited the highest antioxi-
dant activity on DPPH· (65.91% for ME, 61.07% for UAE), ABTS (98.9% for ME,
83.0% for UAE), FRAP (27.0 mg TE/g latex for ME, 24.9 mg TE/g latex for UAE).
Elahe Maghsoudlou et al. (2017) used pulp and skin extracts from two fig culti-
vars (Sabz and Siyah) to enhance the oxidative stability of canola oil. The skin
extract from the Siyah cultivar enhanced the oxidative stability, and its effect was
comparable with a synthetic antioxidant, TBHQ. Therefore, the authors declared
that the Siyah skin extract via subcritical water extraction could be used as a natural
antioxidant in the food system.

5.2 Antimicrobial Activity

There is an increasing interest in plants due to their resisting attributes to antimicro-

bial drugs and the adverse effects of synthetic agents. For this reason, the search for
new antimicrobial compounds, such as phenolic compounds obtained from different
plants, is inevitable (Mahmoudi et al., 2016). Some studies regarding the antimicro-
bial effect of fig, mainly originating from its phenolic compounds, are presented in
Table 17.7. Jeong et al. (2009) reported that MeOH extract of Ficus carica leaves
was inhibitory against some oral bacteria such as S. gordonii, S. anginosus, P. inter-
media, A. actinomycetemcomitans, and P. gingivalis. It was suggested that the flavo-
noids in Ficus carica might be related to the antimicrobial effect observed in this
study, as some flavonoid compounds had intense antimicrobial activity against oral
bacteria. Mahmoudi et al. (2016) extracted phenolic compounds from ten Ficus
carica L. leaves by the soxhlet method. They reported that the tested fig leaves were
good sources of polyphenol and flavonoid compounds and had antimicrobial effects
against Gram-positive and Gram-negative bacteria. Lazreg-Aref et al. (2012) deter-
mined that hexane extract contained high levels of coumarin and reported that pure
coumarins had a strong antimicrobial effect against S. saprophyticus, S. aureus,
E. coli, S. epidermidis, E. cloacae, E. faecalis, and P. aeruginosa. Soni et al. (2014)
evaluated the antibacterial activity of fig extract against Proteus mirabilis and
Bacillus subtilis. Fig extracts exhibited promising antimicrobial activity at 100 mg/
ml concentration against Proteus mirabilis (18.5 mm) and Bacillus subtilis (16 mm).
388 M. Kiralan et al.

Table 17.7 Studies on the antimicrobial activities of fig

Part
used Bioassay Bioactivity References
Leaves Broth dilution The MeOH extract showed strong antibacterial Jeong et al.
activity against oral bacteria. (2009)
Leaves Disc diffusion, The extract of fig leaves showed bactericidal activity Mahmoudi
Macrodilution against G(+) and G(−) bacteria and moderate et al. (2016)
broth antifungal activity.
Leaves Agar well Aqueous extract was active against G(+) bacteria Al Askari
diffusion more than G(−) bacteria but not active against yeast. et al. (2013)
The ethanol extracts presented high antimicrobial
properties against G(+) and G(−).
Latex Disc diffusion, Ethyl acetate extract inhibited the growth of five Aref et al.
Micro-well bacterial species. (2010)
dilution
Latex Disc diffusion, The methanol extract inhibited the growth of Lee and Cha
Micro-dilution Staphylococcus aureus. (2010)
Dried Well dilution The dried fig extract showed inhibition against Soni et al.
fig Bacillus subtilis and Proteus mirabilis. (2014)

5.3 Anticancer Activity

Plants have been used for medicinal purposes since ancient times and are still used
in alternative medicine today. Studies have shown that phytochemicals in plants
have many biological activities such as antioxidant, anticancer, antibacterial, and
anti-inflammatory activities (Mawa et al., 2013). For example, anticancer activity is
agents’ ability with anticancer properties to reverse, stop or slow down the progres-
sion of carcinogens (Chanda & Nagani, 2013). For this reason, plants used in the
treatment of many diseases also play an essential role in treating cancer, which
causes the most deaths in the world in terms of their anticancer properties. Therefore,
many anticancer drugs have been derived from plant-based sources.
In addition to herbs and plant-based drugs, fruits and vegetables also act as anti-
carcinogenic agents to prevent cancer. Fig is an excellent source of phytochemicals
and anticancer medicine regarding seeds, leaves, and roots (Mawa et al., 2013). The
fig fruit that does not contain fat has a rich content of amino acids, minerals, vita-
mins, phenolic compounds, and fiber. No substantial loss was reported in terms of
phenolic compounds and phytochemicals in dried figs (Gençdağ et al., 2021), and it
can maintain its anticancer properties for a long time (Hashemi et al., 2011). Many
studies have been conducted on the anticancer properties of figs and their effects on
health. For example, the effects of fig leaf extract on four types of human cancer
cells: colon cancer, laryngeal carcinoma, hepatocellular carcinoma, and breast can-
cer were investigated and it was stated that this extract had a great potential inhibi-
tory effect, especially on laryngeal carcinoma and hepatocellular carcinoma
(Abdel-Rahman et al., 2021).
Similarly, the extract obtained from the fig fruit was reported to be effective on
breast cancer. Moreover, the fruit extract was also reported to have inhibitory prop-
erties on breast cancer (Jasmine et al., 2015). It was also reported that silver
17 Composition and Functional Properties of Fig (Ficus carica) Phenolics 389

nanoparticles synthesized using fig extract showed an anti-carcinogenic effect

against breast cancer cells (Jacob et al., 2017).
Purnamasari et al. (2019) studied the growth inhibition in liver cancer cells using
the fruit and leaf extracts obtained from fig. According to their results, it was deter-
mined that the extract obtained from the leaf inhibited the growth of liver cancer
cells more than that of the fruit extract.
The anticancer properties of figs are based on the bioactive compounds and anti-
oxidant properties that it contains (Soltana et al., 2019). Antioxidants have the pro-
tective mechanisms to alleviate cancers by improving the immune system. Therefore,
the fig is a natural remedy that helps treat various ailments related to the immune
system, such as gastrointestinal, respiratory, and cardiovascular disorders and can-
cer (Mawa et al., 2013). Furthermore, Jasmine et al. (2015) highlighted a strong
correlation between the antioxidant activity and the anticancer activity of fig.
On the other hand, the factors that inhibit the growth of cancer cells could be due
to other bioactive compounds that are naturally present in fig. It was observed that
fig extracts, which do not show anti-proliferative effects in normal cell lines, cause
cytotoxicity in breast cancer cells (Uz et al., 2015). Therefore, it can be stated that
the bioactive compounds found in the fig fruits exhibit anti-proliferative properties
acting directly on the cancer cells. Moreover, the inhibition of the growth of cancer
cells might occur due to the individual bioactive compounds contained in the fig and
their interaction with each other (Boyacıoğlu et al., 2021). Studies in the literature
support using figs for therapeutic purposes, especially cancer, due to their phyto-
chemical content.

6 Conclusion

This chapter focused on phenolic compounds‘presence and biological activities in

Ficus carica fruits. More than 800 species with 2000 varieties of the Ficus genus are
defined by different researchers. In this chapter, a thorough review has been per-
formed by considering the research findings up to this time. In this respect, phenolic
constituents, i.e., phenolic acids, flavonoids, and anthocyanins isolated by different
studies for various species, were compiled and presented. It can be concluded that
the phenolic composition of fig fruits is highly dependent upon cultivar, harvesting
practices, soil type, geography, and the extraction and analysis methods used. The
novel extraction techniques such as supercritical fluid extraction, microwave-
assisted extraction, and ultrasound-assisted extraction have simultaneous advan-
tages in the extraction of fig phenolics by being more efficient and
environmental-friendly. The figs’ flesh and peel, leaf, and seed parts contain many
phenolic compounds with potent antioxidant, anti-microbial and anticarcinogenic
activities. Therefore, industrial by-products or wastes of fig fruits should also be
evaluated and valorized. Although there are several studies regarding the biological
activities of fig fruits, future investigations are needed on in vitro and in vivo studies
to reveal the further beneficial properties of phenolic compositions of fig fruits.
390 M. Kiralan et al.

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Chapter 18
Phenolic Compounds of Fresh and Dried
Figs: Characterization and Health Benefits

Aicha Debib and Soumaya Menadi

Abbreviations

ABTS 2,2-azinobis-(3-ethylbenzothiazoline-6-
sulphonic acid)
BHT 1% tert-butyl-4-methyl-phenol
DM dry matter
DPPH· 2,2-diphenyl-1-picrylhydrazyl
DW Dry weight
FRAP Ferric ion reducing antioxidant power
FW Fresh weight
GC-MS Gas chromatography-mass spectrometric
HPLC-DAD High-performance liquid chromatography cou-
pled to diode array detection
MDA Malondialdehyd
MS Mass spectroscopy
ORAC Oxygen radical absorbance capacity assay.
ROS Reactive oxygen species
TFC Total flavonoids content
TPC Total phenolics content
UHPLC-QTOF-MS and MS/MS Ultra-high-performance liquid chromatography
coupled to quadrupole-time-of-flight mass
spectrometry
UV-Vis Ultraviolet-visible

A. Debib (*)
Management and Valuation of Agricultural and Aquatic Ecosystems Laboratory,
Morsli Abdellah Tipaza University, Tipaza, Algeria
S. Menadi
Department of Molecular Biology and Genetics, Tokat Gaziosmanpasa University,
Tokat, Turkey
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 395
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_18
396 A. Debib and S. Menadi

1 Introduction

The idea that food could have medicinal properties is not new. Hippocrates, the
father of medicine, proposed the concept “Let food be thy medicine and medicine
be thy food” 2500 years ago. Fruits and vegetables are primary food sources provid-
ing essential nutrients for sustaining life. They also contain a variety of phytochemi-
cals, such as phenolics and flavonoids, which provide important health benefits
(Liu, 2013; Samtiya et al., 2021). Hence, regular consumption of fruits and vegeta-
bles is associated with reduced risks of chronic diseases, such as cancers and cardio-
vascular disease (Dragsted et al., 1993; Jideani et al., 2021; Kaur & Kapoor, 2001).
Recently, attention has focused more on assessing the distribution of biologically
active compounds among different cultivars (Elkordy et al., 2021). In addition,
increasing recent interest in nutraceuticals and functional foods has led plant breed-
ers to initiate the selection of crops with higher-than-normal phenolic antioxidant
contents (Hernandez-Ledesma & Martinez-Villaluenga, 2021). Figs fruits are rich
sources of bioactive compounds such as phenolics, anthocyanins, carotenoids, and
minerals (Cano-Lamadrid & Artés-Hernández, 2021; Meziant et al., 2021). They
are infructescences of the fig (Ficus carica L.), a deciduous tree belonging to the
Moraceae family (Fig. 18.1). Based on archeological findings, figs were probably
one of the first domestically used plants, ca. 12,000 years ago (Abbo et al., 2015;
Kislev et al., 2006). It is an important crop worldwide for both fresh and dry con-
sumption. In 2020, the total world production of figs was 1,260,000 metric tonnes
(FAOSTAT, 2020).
The countries along the Mediterranean coast produce most of the world’s figs. In
these countries, figs are important constituents of the Mediterranean diet, which is
considered one of the healthiest nutritional habits and associated with

Fig. 18.1 Fig fruit anatomy

18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 397

longevity(Veberic & Mikulic-Petkovsek, 2016). Bioactive compounds of figs fruits,

their characterization and utilization in functional foods, and clinical assessment for
human health are among the major targets of contemporary research. There is cur-
rently heightened interest focused on figs as major sources of antioxidants that may
protect against a wide variety of human diseases (Caliskan, 2015). Previously, sev-
eral studies have reported on the health-promoting potential of figs due to the pres-
ence of high concentrations of polyphenols, specifically in dried figs (Arvaniti et al.,
2019; Chang et al., 2016; Debib et al., 2014; dos Anjos Cruz et al., 2021). According
to our knowledge, the detailed information about the health-promoting components
of figs is needed to better understand their use as functional foods and as ingredients
in pharmaceuticals, nutraceuticals and in medicine. Thus, in this review we presents
the main phenolic compounds found in fresh and dried figs from different countries,
describes the analytical methods used for their identification and discuss theirs bio-
logical activities.

2 Research Methods

The search for articles (original or review) was performed using PubMed, Web of
Science, Google Scholar, and Scopus electronic databases until March 2022. The
main keywords used were: “phenolic components”; “Fig fruit”; “Ficus carica”;
“Biological activities”; “antioxidant activity”; “antiviral activity”; “hepatoprotec-
tive effects,”; “anti-microbial activity” and “health benefits”, among others. All the
included references were manually selected and reviewed by the authors.

3 Fig Phytochemicals

3.1 Extraction and Quantification Processes

Extraction is a crucial step in isolating phenolic chemicals, as well as later iden-

tifying and quantifying them (Konak et al., 2015). Various extraction methods
have been used, ranging from traditional solvent extraction using water or organic
solvents to alternative techniques that use greener solvents or intensification tools
to improve yields and rates (Chemat et al., 2019). From literature data, it is well
known that the solvent and temperature are the process parameters that have the
largest impact on the efficiency of extracting bioactive chemicals from plant mate-
rial (Bucic-Kojic et al., 2011). Indeed, the selection of the solvent is one of the most
important steps in extraction. Methanol, ethanol, and their mixtures in water, as
well as the acetone, petroleum ether, and dimethylformamide, are the most com-
monly used solvents in the extraction of fig fruit phenolic compounds (Naczk &
Shahidi, 2006; Nakilcioğlu-Taş & Ötleş, 2021). Moreover, ethanol and water are
398 A. Debib and S. Menadi

much safer and more suitable for the food and pharmaceutical industry than the
other organic solvents from a toxicological point of view. According to Bucic-Kojic
et al. (2011) and Debib et al. (2014), the fig variety and extraction process con-
ditions significantly affected the content of phenolic compounds in the fig fruits.
These two studies reported that the best extraction efficiency was obtained with
80% (v/v) aqueous ethanol solvent. The presence of the highest total phenolic
content in aqueous solutions could be attributed to the increase in the polarity of
the solvents by the addition of water. On the other hand, several studies reported
that ultrasonic extraction generally improves extraction efficiency (Bimakr et al.,
2017; Júnior et al., 2021; Vardanega et al., 2014). Thongson et al. reported that the
ultrasonic extraction method only took 5 min to obtain the bioactive components
of a medicinal plant (Thongson et al., 2004). Although their structural complexity
and diversity hamper the quantitative determination of polyphenols, several meth-
ods have been used to determine polyphenols in fig fruit extracts. The colorimetric
Folin-Ciocalteu essay is the most basic method used; however, modern analytical
tools have helped to reveal the highly diverse and complicated chemical structure
of phenolic fig compounds, usually using advanced and combined techniques such
as GC-MS, HPLC-DAD-ESI/MS, RP-HPLC-PDA, and LC-ESI-MS/MS (Tenório
et al., 2022). Many studies have discussed the use of the Folin-Ciocalteau reagent to
determine polyphenols and the general or a specific value of the method (Blainski
et al., 2013; Margraf et al., 2015; Míguez et al., 2022). In alkaline solutions, this
reagent decomposes quickly, necessitating the use of a large amount of the reagent
to complete the reaction. Precipitation and severe turbidity can ensue from this
excess, making spectrophotometric analysis unfeasible. Folin and Ciocalteu solved
this problem by include lithium salts in the reagent، which prevented turbidity. The
reaction generally provides accurate and specific data for several groups of phenolic
compounds, because many compounds change color differently due to differences
in unit mass and reaction kinetics (Blainski et al., 2013).

3.2 Phenolic Composition of Fig Fruit

The term phenolic compound involves a wide range of various solvent-soluble com-
pounds, having at least one aromatic ring with one or more hydroxyl groups
attached. Based on their structural skeletons, these compounds are broadly divided
into simple phenolics, phenolic acids, coumarins, flavonoids, stilbenes, tannins, lig-
nans, and lignins (Miklavčič Višnjevec & Schwarzkopf, 2020; Vuolo et al., 2019).
During the last decade, phytochemical studies on dried and fresh figs revealed the
presence of numerous bioactive compounds such as phenolic compounds, phytos-
terols, organic acids, anthocyanin composition, triterpenoids, coumarins, and vola-
tile compounds such as hydrocarbons, and a few other classes of secondary
metabolites. However, the fruit peel presented the highest value of phenolic com-
pounds. Indeed, the amount of fresh figs was higher than dried figs. In literature, the
reason for this was explained as the demolition of some polyphenols of the fresh figs
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 399

during the drying process or the transformation to non-antioxidant forms

(Nakilcioğlu & Hışıl, 2013). The majority of studies on phenolic compounds in figs
originates from Algeria and Tunisia (Ammar et al., 2015; Bachir Bey et al., 2017;
Aicha Debib et al., 2016, 2014; Faleh et al., 2012; Kehal et al., 2021; Khadhraoui
et al., 2019; Mahmoudi et al., 2016; Meziant et al., 2021; Soltana et al., 2018) and
Turkey (Çalişkan & Polat, 2011; Ercisli et al., 2012; Kamiloglu & Capanoglu, 2013;
Konak et al., 2015; Nakilcioğlu-Taş & Ötleş, 2021; Nakilcioğlu & Hışıl, 2013;
Yemiş, Bakkalbaşı, & Artık, 2012). While there is less data available from Spain
(Dueñas et al., 2008; Vallejo et al., 2012; Viuda-Martos et al., 2015; Wojdyło et al.,
2016), Slovenia (Slatnar et al., 2011; Veberic et al., 2008), Italy (Del Caro & Piga,
2008; Piga et al., 2005), Croatia (Bucic-Kojic et al., 2011; Jokić et al., 2014), China
(Feng et al., 2015; Qin et al., 2015), Iran (Maghsoudlou et al., 2017), United States
(Crisosto et al., 2010; Kehal et al., 2021; Vinson et al., 2005) and Portugal (Oliveira
et al., 2009; Palmeira et al., 2019,

3.2.1 Phenolic Acids

Phenolic acids consist of two subgroups, hydroxybenzoic and hydroxycinnamic

acids. Hydroxybenzoic acids include gallic, p-hydroxybenzoic, protocatechuic,
vanillic, and syringic acids having a C6-C1 structure, whereas hydroxycinnamic are
aromatic compounds with a three-carbon side chain (C6–C3), with caffeic, ferulic,
p-coumaric, ellagic and chlorogenic acids. The main phenolic acid compounds
identified in the Ficus carica fruits are described in Table 18.1 According to Arvaniti
et al. (2019), the most predominant phenolic acids found in dried figs are gallic acid
up to 5.04 mg/100 g Dry Matter (DM) and chlorogenic acid up to 32.4 mg/100 g
DM, while the same target compounds in whole fresh figs have been determined at
concentrations up to 6.98 mg/100 g DM (gallic acid) and 28.8 mg/100 g DM (chlo-
rogenic acid) (Arvaniti et al., 2019).
Apart from the aforementioned phenolic acids, figs have also been reported to
have the presence of various coumarins such as psoralen, bergapten, marmesin,
umbelliferone, and angelicin (Veberic & Mikulic-Petkovsek, 2016). In addition,
methanolic extracts of fig were shown to possess good psoralen levels, methoxyp-
soralen, and bergapten (Oliveira et al., 2010; Ammar et al., 2015).

3.2.2 Flavonoids

The generic structure of flavonoids consists of two aromatic rings (A and B rings)
linked by three carbons, usually in an oxygenated heterocycle ring called a C ring
(Fig. 18.2). Based on differences in the heterocycle C ring, flavonoids are catego-
rized as flavonols (quercetin, kaempferol, and myricetin), flavones (luteolin, and
apigenin), flavanols (catechins, epicatechin, epigallocatechin, and epicatechin gal-
late), flavanones (naringenin), anthocyanidins, or isoflavonoids (genistein, daid-
zein, dihydrodaidzein, and equol) (Yang & Xiao, 2013). Figs flavonoids are
400 A. Debib and S. Menadi

Table 18.1 Phenolics acids compounds from Ficus carica fruits

Analytical
Structure Compound Sample preparation method Reference
p-hydroxyben Methanolic extract HPLC- Aicha Debib et al.
zoic acid R, DAD- ESI/ (2016)
R2 = H, R1, MS
R3 = OH
Gallic acid R, Aqueous extract HPLC Rtibi et al. (2018),
R1, R2, Hydromethanolic HPLC-DAD Alamgeer et al.
R3 = OH extract HPLC/ (2017), De Masi
Hydromethanolic UV-DAD et al. (2015), and
extract LC-ESI-MS Khadhraoui et al.
Pressurized liquid (2019)
extraction and
solid-phase
extraction
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Ultrasonic HPLC-PDA Veberic et al. (2008)
extraction
Vanillic acid Hydroethanolic RP-UHPLC- Belguith-Hadriche
R2 = OCH3 extract DAD- et al. (2016)
QTOF-MS
and MS/ MS
Hydromethanolic HPLC-DAD Alamgeer et al.
extract (2017)
Methanolic extract HPLC- Aicha Debib et al.
DAD- ESI/ (2016)
MS
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Syringic acid R, Aqueous extract HPLC Rtibi et al. (2018)
R2 = OCH3,
R1 = OH
Hydromethanolic HPLC-DAD Alamgeer et al.
extract (2017)
Aqueous extract HPLC Nakilcioğlu and
Hışıl, (2013)
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Ultrasonic HPLC-PDA Veberic et al. (2008)
extraction
Protocatechuic Hydromethanolic HPLC/ De Masi et al.
acid R = H, R1, extract UV-DAD (2015)
R2, R3 = OH Pressurized liquid LC-ESI-MS Khadhraoui et al.
extraction and (2019)
solid-phase
extraction
Cinnamic acid Pressurized liquid LC-ESI-MS Khadhraoui et al.
R, R1 = H extraction and (2019)
solid-phase
extraction

Ethyl acetate extract GC-MS Ahmad et al. (2016)

(continued)
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 401

Table 18.1 (continued)

Analytical
Structure Compound Sample preparation method Reference
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Methanolic extract HPLC- Aicha Debib et al.
DAD- ESI/ (2016)
MS
p-coumaric acid Methanolic extract HPLC- Aicha Debib et al.
R = OH, R1 = H DAD- ESI/ (2016)
MS
Hydromethanolic HPLC Kamiloglu and
ultrasonic extraction Capanoglu (2013)
m-coumaric acid Hydromethanolic HPLC-DAD Alamgeer et al.
R = H, R1 = OH extract (2017)
Caffeic acid Ethyl acetate extract GC-MS Ahmad et al. (2016)
R, R1 = OH
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Caffeic acid Hydroethanolic LC-DAD- Palmeira et al.
hexoside extracts ESI/MSn (2019)
R = OH,
R1 = O-hexoside
Ferulic acid Ethyl acetate extract GC-MS Ahmad et al. (2016)
R = OH,
R1 = OCH3
Acetone extract HPLC/DAD Yemiş et al. (2012)
and HPLC/
MS
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Hydroethanolic RP-UHPLC- Belguith-Hadriche
extract DAD- et al. (2016)
QTOF-MS
and MS/ MS
Pressurized liquid LC-ESI-MS Khadhraoui et al.
extraction and (2019)
solid-phase
extraction
Hydromethanolic HPLC/ De Masi et al.
extract UV-DAD (2015)
Hydromethanolic HPLC Bachir Bey et al.
extract (2017)
Acid water (pH 2 HPLC-DAD Faleh et al. (2012)
with HCl)
Chlorogenic Hydroethanolic LC-DAD- Palmeira et al.
acid R = OH extracts ESI/MSn (2019)
Ultrasonic HPLC-PDA Veberic et al. (2008)
extraction
(continued)
402 A. Debib and S. Menadi

Table 18.1 (continued)

Analytical
Structure Compound Sample preparation method Reference
Solid-phase UHPLC- Yemiş et al. (2012)
extraction MS-CID
Aqueous extract HPLC Nakilcioğlu and
Hışıl (2013)
Aqueous extract HPLC Rtibi et al. (2018),
Hydromethanolic HPLC/ Alamgeer et al.
extract UV-DAD (2017), De Masi
Hydromethanolic LC-ESI-MS et al. (2015), and
extract Khadhraoui et al.
Pressurized liquid (2019)
extraction and
solid-phase
extraction
Ellagic acid Aqueous extract HPLC Rtibi et al. (2018)
Acetone/ formic HPLC-DAD/ Pereira et al. (2020)
acid extract ESI-MS
Hydromethanolic HPLC Kamiloglu and
ultrasonic extraction Capanoglu (2013)

Psoralen Aqueous extract HPLC/DAD Oliveira et al.

and HPLC/ (2009)
UV
Furanocoumarins Hydroethanolic RP-UHPLC- Belguith-Hadriche
extract DAD- et al. (2016)
QTOF-MS
and MS/ MS
Bergapten Aqueous extract HPLC/DAD Oliveira et al.
and HPLC/ (2009)
UV
Hydroethanolic RP-UHPLC- Belguith-Hadriche
extract DAD- et al. (2016)
QTOF-MS
and MS/ MS
Marmesin Ultrasonic HPLC-PDA Veberic et al. (2008)
extraction

Umbelliferone Ultrasonic HPLC-PDA Veberic et al. (2008)

extraction

Angelicin Ultrasonic HPLC-PDA Veberic et al. (2008)

extraction
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 403

primarily categorized into flavanols, flavonols, flavones, and anthocyanidins

(Table 18.2).

4 Health Benefits of Figs

The positive health effects of fresh and dried figs have been attributed to many fac-
tors, mainly phenolic compounds. In addition, numerous epidemiological data, ani-
mal work, and cell culture evidence support the health benefits of figs in preventing
cardiovascular disease and certain cancers (Alamgeer et al., 2017).

4.1 Antioxidant Activity

From the chemical standpoint, the antioxidants act by neutralizing products of

metabolism, such as free radicals, thus reducing degenerative processes related to
aging phenomena. It is well known that free radicals can cause damage to cells and
their constituents that can lead to the onset and development of chronic degenerative
diseases. In fact, under normal physiological conditions, cellular metabolism pro-
duces reactive oxygen species (ROS) in equilibrium with the antioxidant defense
system. When this homeostasis is interrupted by excess ROS, oxidative stress. In
this situation, ROS can cause lipid peroxidation (e.g., increased malondialdehyde
[MDA]) and oxidative damage to DNA molecules (e.g., cancer), membrane phos-
pholipids, and proteins, which constitute the molecular basis of several diseases
(Khutami, Sumiwi, Khairul Ikram, & Muchtaridi, 2022; Niemann et al., 2017). In

Fig. 18.2 Structure of main classes of figs flavonoids

404 A. Debib and S. Menadi

Table 18.2 Flavonoids compounds from Ficus carica fruits

Analytical
Flavonoids class Compound Sample preparation method Reference
Flavones Luteonil-8- Methanolic extract HPLC-MS Slatnar et al.
Cglucoside (2011)
Methanolic extract HPLC-DAD- Aicha Debib
ESI/MS et al. (2016)
Luteolin-6-C- Pressurized liquid LC-ESI-MS Khadhraoui
Apentose extraction and et al. (2019)
Apigeniun solid-phase
extraction
Aqueous extract HPLC Rtibi et al.
(2018)
Apigenin-2′′ Hydroethanolic LC-DAD- Palmeira et al.
-O-rhamnose-C- extracts ESI/MSn (2019)
acetylhexoside
Orientin Hydroethanolic RP-UHPLC- Belguith-
extract DAD- Hadriche et al.
QTOF-MS (2016)
and MS/ MS
Isorientin Hydroethanolic RP-UHPLC- Belguith-
extract DAD- Hadriche et al.
QTOF-MS (2016)
and MS/ MS
(continued)
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 405

Table 18.2 (continued)

Analytical
Flavonoids class Compound Sample preparation method Reference
Flavonols Rutin Acetone/ formic acid HPLC-DAD/ Pereira et al.
extract ESI-MS (2020)
Aqueous extract HPLC Rtibi et al.
(2018)
Hydroethanolic LC-DAD- Palmeira et al.
extracts ESI/MSn (2019)
Hydroethanolic RP-UHPLC- Belguith-
extract DAD- Hadriche et al.
QTOF-MS (2016)
and MS/ MS
Hydromethanolic HPLC/ De Masi et al.
extract UV-DAD (2015)
Quercetin Hydromethanolic HPLC/ De Masi et al.
extract UV-DAD (2015)
Hydroethanolic RP-UHPLC- Belguith-
extract DAD- Hadriche et al.
QTOF-MS (2016)
and MS/ MS
Hydromethanolic HPLC-DAD Alamgeer et al.
extract (2017)
Methanolic extract UHPLC- Soltana et al.
QTOF-MS/ (2018)
Methanolic extract HPLC-DAD- Aicha Debib
ESI/MS et al. (2016)
Quercetin-3-O- Hydroethanolic RP-UHPLC- Belguith-
(malonyl)- extract DAD- Hadriche et al.
glucoside QTOF-MS (2016)
and MS/ MS
Methanolic extract UHPLC- Soltana et al.
QTOF-MS/ (2018)
Quercetin-3-O- Acetone/ formic acid HPLC-DAD/ Pereira et al.
(acetyl)- glucoside extract ESI-MS (2020)
Isoquercitrin Methanolic extract HPLC-DAD- Aicha Debib
ESI/MS et al. (2016)
Methanolic extract UHPLC- Soltana et al.
QTOF-MS (2018)
Hydroethanolic RP-UHPLC- Belguith-
extract DAD- Hadriche et al.
QTOF-MS (2016)
and MS/ MS
Quercetin-3- Hydromethanolic HPLC Kamiloglu and
Orutinoside ultrasonic extraction Capanoglu
(2013)
Kaempeferol-3- Methanolic extract HPLC-MS Slatnar et al.
O-glucoside (2011)
(continued)
406 A. Debib and S. Menadi

Table 18.2 (continued)

Analytical
Flavonoids class Compound Sample preparation method Reference
Flavanols Catechin Hydromethanolic HPLC Kamiloglu and
ultrasonic extraction Capanoglu
(2013)
Methanolic extract HPLC-MS Slatnar et al.
(2011)
Hydromethanolic HPLC Bachir Bey
extract et al. (2017)
Epicatechin Acetone/ formic acid HPLC-DAD/ Pereira et al.
extract ESI-MS (2020)
Aqueous extract HPLC Rtibi et al.
(2018)
Pelargonidin-3-O- Acetone/ formic acid HPLC-DAD/ Pereira et al.
rutinoside extract ESI-MS (2020)
Anthocyanidins Cyanidin-3-O- Acetone/ formic acid HPLC-DAD/ Pereira et al.
rutinoside extract ESI-MS (2020)
Methanolic extract HPLC-MS Slatnar et al.
(2011)
Hydromethanolic HPLC Kamiloglu and
ultrasonic extraction Capanoglu
(2013)
Hydromethanolic HPLC Bachir Bey
extract et al. (2017)
Cyanidin-3-O- Hydromethanolic HPLC Bachir Bey
glucoside extract et al. (2017)
Solid-phase UHPLC-MS- Yemiş et al.
extraction CID (2012)
Hydromethanolic HPLC Kamiloglu and
ultrasonic extraction Capanoglu
(2013)
Cyanidin-3,5- Solid-phase UHPLC-MS- Yemiş et al.
Diglucoside extraction CID (2012)
Peonidin Methanolic extract HPLC-DAD- Dueñas et al.
3-O-glucoside ESI/MS (2008)
Pelargonidin Methanolic extract HPLC-DAD- Dueñas et al.
3-O-glucoside ESI/MS (2008)
Pelargonidin 3- Acetone/ formic acid HPLC-DAD/ Pereira et al.
O-rutinosid extract ESI-MS (2020)

addition, antioxidants inhibit the action of enzymes that catalyze the detoxification
pathway in the body upon exposure to mutagens. Based on literature data, several
studies reported that flavonoids have an antioxidant action that attenuates ROS by
enhancing antioxidant enzymes or by inhibiting enzymes that employ a pro-oxidant
effect on oxidative stress (e.g., NADPH oxidase, xanthine oxidase, and catalase)
(Yan et al., 2013; Zhang et al., 2022).
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 407

The antioxidant capacity of different components of fresh figs (skin, pulp, entire
fruit) and dried figs have been intensely studied using a variety of antioxidant
assays. Some of the methods that have been widely used in selected fig varieties
include the 1,1-diphenyl-2-picrylhydrazyl radical assay (DPPH·), 2,2-azino-di-
(3-ethylbenzothialozine-sulphonic acid) assay (ABTS), ferric ion reducing antioxi-
dant power assay (FRAP), cupric ion reducing capability assay (CUPRAC), and
oxygen radical absorbance capacity assay (ORAC). According to the most pub-
lished results, the antioxidant capacity of figs has been highly correlated with the
amount of phenolic compounds (Ouchemoukh et al., 2012; Vinson et al., 2005;
Wojdyło et al., 2016). Furthermore, according to khadhraoui et al. (2017), there is a
strong correlation between skin color, total phenolic content, and antioxidant activ-
ity. Thus, figs from dark-colored cultivars exhibited a higher antioxidant capacity
than those from light-colored cultivars, with 3–4 fold differences between cultivars.
These findings were confirmed by several studies (Faleh et al., 2012; Konak et al.,
2015; Pereira et al., 2020). Regarding the impact of drying on the antioxidant capac-
ity, Chang et al. (2016) reported that the sun-drying process had a positive effect on
antioxidant activity, which increases the antioxidant activity of dried figs compared
to that of fresh figs (Chang et al., 2016). While in other studies, it has been reported
that the sun-drying procedure reduced the antioxidant capacity of figs (Bachir Bey
et al., 2017; Kamiloglu & Capanoglu, 2015) (Fig. 18.3).

4.2 Antimicrobial Activity

Antibacterial activity has been demonstrated in several varieties of Ficus carica

fruit worldwide. Debib et al. (2014) reported that four different extracts of two
Algerian figs varieties (Azendjar and Taamriout) showed antimicrobial effects
against gram-positive bacteria such as Staphylococcus aureus (ATCC 25923),
Enterococcus faecalis (ATCC 29212) and Bacillus subtilis subsp. spizizenii (ATCC
6633) and six gram-negative bacteria, Pseudomonas aeruginosa (ATCC 27853),
P. aeruginosa (ATCC 10145), Escherichia coli (ATCC 25922), Enterobacter cloa-
cae (ATCC 13047), Salmonella enterica subsp. heindelberg (ATCC 8326),
Klebsiella pneumoniae (clinical isolated), and one yeast, Candida albicans (clinical
isolated). According to these authors, this effect could be attributable to these
varieties‘high flavonoid and tannin levels. Their ability to combine with extracellu-
lar and soluble proteins and bacterial cell walls is most probably the cause of their
activity. Indeed, microbial membranes may be disrupted by lipophilic flavonoids. In
another study, the intense antibacterial activity of the figs methanolic extract was
found against Staphylococcus aureus, Bacillus subtilis, Bacillus cereus, Listeria
monocytogenes, Salmonella typhimurium, and Escherichia coli, mainly against
gram-positive bacteria (Shahbazi, 2017). The peel and pulp that have been separated
from the F. carica fruit (ethanol/water extract, 80:20 v/v) showed similar results in
most bacterial strains tested. The best results were obtained against gram-positive
408 A. Debib and S. Menadi

Fig. 18.3 Imbalance between reactive oxygen species (ROS), a possible antioxidant effect of
phenolic compounds of figs

bacteria, especially methicillin-sensitive S. aureus, with a MIC of 2.5 mg/mL

(Palmeira et al., 2019).
The oily dried figs macerates extracts possessed significantly higher antimicro-
bial activity than the oily extracts against Staphylococcus aureus, Enterococcus fae-
calis, Bacillus subtilis Pseudomonas aeruginosa, Escherichia coli, Enterobacter
cloacae, Salmonella enterica. Antimicrobial activity results of this investigation
validate the use of olive oily macerates of dried figs by Algerian people to fight
infectious diseases (Debib et al., 2018).

4.3 Antidiabetic Effect

It has been revealed that long-term use of anti-diabetic medications is linked to

several toxicities harmful to human health. Anti-diabetic fruits and vegetables could
be a valuable tool in developing safer and more effective oral hypoglycemic treat-
ments; similarly, the discovery of anti-diabetic therapies has turned to natural plant
sources with few adverse effects (Gothai et al., 2016). Different extracts of fruits
and leaves showed potent inhibitory activity against α-amylase and α-glucosidase
enzymes (Wojdyło et al., 2016). According to El-Shobaki et al. (2010), fruits and
leaves of F. carica showed intense antidiabetic activity in alloxon-induced diabetic
rats by increasing antioxidant levels. In other studies, the methanol extract of fruits,
aqueous leaves, and decoction obtained from the leaves also controlled the diabetic
complications in streptozotocin-induced diabetic rats (Belguith-Hadriche et al.,
2016; Mopuri & Islam, 2016). Several in vivo investigations have proposed
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 409

mechanisms of action (Meziant et al., 2021; Perez et al., 2003). Pèrez et al. (2003)
and Belguith-Hadriche et al. (2017) demonstrated that Ficus carica extracts have
positive effects on insulin secretion and blood glucose levels (Perez et al., 2003).
Song et al. (2002) reported flavonoids as hypoglycaemic agents because they
improved glucose metabolism and reduced oxidative stress in diabetic rats (Song
et al., 2002). When the diabetic rats were given glucose with quercetin, hyperglyce-
mia significantly decreased compared with glucose administered alone, and flavo-
noids modulated glucose transport via their intestinal transporter. The same results
were obtained in human subjects, F. carica fruit extract standardized in abscisic acid
was added to a glucose beverage, reducing the postprandial glycemic and insulin-
emic values in healthy adult subjects when compared to the normal drink (Atkinson
et al., 2019).
The hydromethanolic extract of F. carica has also been shown to have anti-
obesity properties in recent in vivo investigations. The results were mainly evi-
denced by the reduction in the animals’ body mass and the serum levels of
low-density lipoprotein (LDL) (Arafa et al., 2020).

4.4 Anticancer Activity

The methanol extract of F. carica at a dose of 1000 μg/mL revealed low anti-cancer
activity through apoptosis (0.44%) and necrosis (2.26%) of liver cancer cells
(Huh7it). The positive control (doxorubicin, 10 μg/mL) reached 65.6 and 34.1%,
respectively (Purnamasari et al., 2019). Sixteen prenylated isoflavone derivatives
isolated from the ethanol extract presented an antiproliferative effect against human
cancer cell lines, HL-60 (Liu et al., 2019). In another study, cytotoxicity of fruit and
leaf extracts, as well as the latex on HeLa cell line, was examined, and results
showed that latex and different extracts could reduce the viability of HeLa cells at
concentrations as low as 2 μg/mL in a dose-dependent manner (Khodarahmi
et al., 2011).

4.5 Anti-Inflammatory Activity

Inflammation is a natural response of the body against microorganisms and toxic

materials. Various natural products have proven their role as anti-inflammatory
agents without severe side effects. Debib et al. (2016) reported that the methanolic-
dried figs fruit extract at a dose of 38.07 mg/kg/day enhanced the protection of
CCl4-induced oxidative stress and hepatotoxicity in rats. According to these studies,
these hepatoprotective effects could be attributed to the inhibitory activity of flavo-
noids on free radical production since exogenous antioxidants may counteract the
damaging effects of oxidative stress, cooperating with natural systems like glutathi-
one tocopherol or protective enzymes.
410 A. Debib and S. Menadi

4.6 Effect on Alzheimer’s Disease

Subash et al. (2016) have studied the effect of dietary supplementation with 4% figs
grown in Oman on the memory, anxiety, and learning skills in APPsw/Tg2576 (Tg
mice) mice model for Alzheimer’s disease. The authors have assessed spatial mem-
ory and learning ability, psychomotor coordination, and anxiety-related behavior in
Tg and wild-type mice at the age of 4 months and after 15 months using the Morris
water maze test, rota-rod test, elevated plus maze test, and open-field test. Results of
these study revealed that Tg mice fed a control diet without figs showed significant
memory deficits, increased anxiety-related behavior, and severe impairment in spa-
tial, position discrimination learning ability, and motor coordination compared to
the wild-type control mice on the same diet, and Tg mice fed on 4% fig diet supple-
mentation for 15 months. These studies suggest that dietary supplementation of figs
may help improve cognitive and behavioral deficits in Alzheimer’s disease (Subash
et al., 2016). Another study investigated the impact of dietary supplements, includ-
ing pomegranate, figs, or dates, on inflammatory cytokines suppression. The
researchers discovered that using the food supplements reduced inflammatory cyto-
kines and eotaxin activity (Essa et al., 2015).

4.7 Antispasmodic Activity

The antispasmodic efficacy of an aqueous-ethanolic extract of dried figs was inves-

tigated. The research revealed the presence of spasmolytic activity in the ripe dried
fruit, which could be mediated by the activation of K + ATP channels (Gilani
et al., 2008).

4.8 Anti-Osteoporotic Potential

Another study found that in RANKL-stimulated RAW264.7 cells and bone marrow-
derived macrophages, the hexane soluble extract of the fig is a potent inhibitor of
osteoclastogenesis (Park et al., 2009). These results corroborate those of Mohammed
et al. (2018), who revealed that 50 mg/kg raw extract solution showed potential
against osteoporosis in ovariectomized rats, comparable to estrogen replacement
therapy (Mohammad et al., 2018; Park et al., 2009).
18 Phenolic Compounds of Fresh and Dried Figs: Characterization and Health Benefits 411

5 Conclusion

The popularity of alternative medicine and natural products is increasing worldwide

due to inexpensive and health-promoting activities. In this vista, Ficus carica Linn
fruit, generally known as figs, is associated with a lowered risk of chronic diseases.
In vitro and in vivo studies have demonstrated that figs phenolic compounds are
involved in a broad spectrum of biological activities, including antioxidant, antimi-
crobial, antidiabetic, anticancer, anti-inflammatory, antispasmodic, and anti-
osteoporotic activities. In addition, fresh and dried fig extract inhibits cancer cell
proliferation and reduces Alzheimer’s disease complications. Pharmacological
studies on the fresh and dried fruit, crude extracts, and isolated components of Ficus
carica provide experimental support for its numerous traditional uses. In this review,
89 papers described the chemical characterization and biological activities of Ficus
carica L. fruit. However, there is a vast scope for establishing a relation between
phytoconstituents and biological activities. The outcome of the future research in
the areas mentioned above will provide convincing support for the future clinical
uses of Ficus carica in modern medicine.

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Chapter 19
Ficus carica L. as a Source of Natural
Bioactive Flavonoids

Leila Meziant and Mostapha Bachir-bey

Abbreviations

C3G Cyanidin-3-Glucoside
C3R Cyanidin-3-Rutinoside
CE Capillary Zone Electrophoresis
DEEs Deep Eutectic Solvents
EAE Enzyme-Assisted Extraction
ILs Ionic-Liquids
IRAE Infrared-Assisted Extraction
LLE Liquid-Liquid Extraction
MAE Microwave-Assisted Extraction
MALDI Matrix Assisted Laser Desorption
MAP Modified Atmosphere Packaging
MCAE MechanoChemical-Assisted Extraction
MS Mass Spectrometry
NIRS Near-Infrared Reflectance Spectroscopy
NMR Nuclear Magnetic Resonance
PAL Phenylalanine Ammonia-Lyase
PEF Pulsed-Electric Field Extraction
PLE Pressurized-Liquid Extraction
RP-HPLC Reverse-Phase High Performance
SFE Supercritical-Fluid Extraction
SPE Solid-Phase Extraction
UAE Ultrasound-Assisted Extraction
UHPLC Ultra-High Performance Liquid Chromatography

L. Meziant · M. Bachir-bey (*)

Laboratory of Applied Biochemistry, Faculty of Natural and Life Sciences, University of
Bejaia, Bejaia, Algeria
e-mail: [email protected], [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 417
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_19
418 L. Meziant and M. Bachir-bey

1 Introduction

Phenolic compounds are by far the most documented phytochemicals in plants.

These heterocyclic molecules (more than 8000 structures found in nature) were at
first considered of relevance just for the plants since phenolic compounds are key
molecules in plant physiology, acting as insect and herbivore repellents, signaling
mediators, and contributing to defense mechanisms against external biotic patho-
gens and non-biotic conditions (Treutter, 2006). However, new applications in food
and health domains have been considered with the discovery of their biological
activities. Such interest in phenolic compounds came from their ability to prevent
and treat different health disorders related to oxidative stress. This high antioxidant
activity is exerted through different mechanisms, including free radical scavenging,
protecting and regenerating other dietary antioxidants, and reducing and chelating
pro-oxidant metal ions (Khan et al., 2019). On the chemical structure basis, pheno-
lic compounds could be divided into nine distinct groups, namely (1) phenolic acids
(hydroxybenzoic acids, hydroxycinnamic acids), (2) flavonoids (anthocyanins, fla-
vanols, flavonols, dihydroflavonols, flavones, isoflavones, flavanones, dihydrochal-
cones), (3) coumarins (coumestans, furanocoumarines), (4) stilbenoids, (5) quinones
(benzoquinones, naphtoquinones, anthraquinones), (6) lignans, (7) xanthones, (8)
phenylpropanoids (benzodioxoles, cucuminoids, hydroxyphenylpropenes) and (9)
phenols (alkylphenols, methoxyphenols) (Murthy & Bapat, 2020). Flavonoids are
the major category of phenolics found in nature, given their scientific importance in
different domains, especially food and health.

2 Structure, Occurrence and Function of Flavonoids

Flavonoids, defined as ubiquitous plant pigments, are liable to yellow, orange, red,
purple, and black colors of the flowers, fruits, grains, and other plant tissues and are
involved in plant growth and defense. Flavonoids are synthesized through the phen-
ylpropanoid pathway (also called the shikimic acid pathway) and subdivided into
six subclasses: flavonols, found in cherries, onion, garlic, and broccoli; flavones
found in artichokes, celery, parsley, and green pepper; flavanones found in oranges,
lemons, grapefruits, and oregano; flavanols found in cocoa, tea, apples, nuts and
blackberries; isoflavones found mainly in soybeans and soy products; and anthocya-
nins found in grapes, blueberries, black currants, red cabbage and chickpeas (Prior,
2003; Raffa et al., 2017). Their chemical structure is based on a C6-C3-C6 carbon
skeleton consisting of two benzene rings (A and B rings) linked by a heterocyclic
pyran ring (C ring). Flavonoids are often hydroxylated in positions 3, 5, 7, 2′, 3′, 4′,
and 5′, and could form methyl ethers or acetyl esters (Kumar & Pandey, 2013). Most
flavonoids are O- or C-linked glycosides, and when glycosides are formed, the gly-
cosidic linkage often occurs in positions 3 or 7 with D-glucose, L-rhamnose, rutin-
ose, galactose, or arabinose (Kumar & Pandey, 2013). Other entities found in nature
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 419

are also considered flavonoids, such as biflavonoids, prenyl-flavonoids, glycosidic

ester flavonoids, flavonolignans, chalcones, and proanthocyanins (Brodowska, 2017).
Flavonoids were first discovered in 1930 by Albert Szent-Györgyi (Nobel Prize
laureate in 1937) as new bioactive substances in Citrus extract classified as “vitamin
P” which stabilized the biological activity of ascorbic acid and cured scurvy. Vitamin
P was later identified as a mixture of hesperidin and eriodictyol-glycoside (flava-
nones) (Perez-Vizcaino & Fraga, 2018). Flavonoids and their natural derivatives are
found in commercialized dietary supplements and numerous pharmaceutical prepa-
rations. They also possess multiple applications (1) in the food industry as flavoring,
sweetening, and coloring agents, (2) in leather production, as tanning agents, (3) in
pigment industry, as dyes or mordant-dyes, (4) in horticulture and flower industry,
as efficient agents against plant viruses, microbial pathogens and wood-destroying
fungi (Brodowska, 2017).
The increasing interest in flavonoids (shown by the growing number of scientific
papers found in this field) could be explained by the fact that these compounds (1)
are highly diversified, widely distributed in the plant kingdom, and easily separable
with chromatographic methods, (2) are natural pigments with an exceptional safety
score and a very wide range of hues (from yellow, orange, red, blue, violet to black)
which make them good substituents of industrial synthetic food colorings, (3) pos-
sess interesting structural and functional properties that could serve as a base for the
development of pharmaceutical drugs, (4) have a long history of medical use for the
treatment of various ailments which make them a dominant class of natural thera-
peutic agents (Ayaz et al., 2019; Havsteen, 2002).

3 Biologically Active Flavonoids

Bioflavonoids identified as biologically active flavonoid compounds constitute an

important part of the human diet. Their intake, as estimated in Australian, European,
and US adult populations, varies considerably from 209 to 1017 mg/day (Peterson
et al., 2015), a variability that can be related to differences in the food supply and
cultural eating patterns between countries (Kent et al., 2018). Flavonoid intake in
the Asian population could be higher (up to 2000 mg/day) because of the high con-
sumption of legumes, soy products, and tea (Kozlowska & Szostak-Wegierek,
2014). Although from the total intake, only 5–10% (mostly monomeric and dimeric
flavonoids) can be absorbed in the small intestine after deglycosylation reactions,
the remaining molecules are metabolized in the colon by the gut microbiota: the
microorganisms can break up the heterocyclic rings, and the formed hydroxylated
phenyl carboxylic acids could be absorbed; flavonoids and the microbial-derived
metabolites reach the plasma after 1 to 2 h after ingestion (Kamiloglu et al., 2020;
Kopustinskiene et al., 2020).
Consumption of flavonoids could improve intestinal barrier function, modulate
the intestinal microbiota and improve microbial dysbiosis in obesity by inhibiting
the growth of pathogenic bacteria and increasing the probiotic genera such as
420 L. Meziant and M. Bachir-bey

Bifidobacterium and Lactobacillus which could improve intestinal health by reduc-

ing endotoxin production, rising the conversion of primary into secondary bile
acids, maintaining intestinal immune homeostasis and promoting nutrients absorp-
tion (Pei et al., 2020). Flavonoid-rich foods are associated with a lower incidence of
cardiovascular diseases, cancer, and all-cause mortality: intake of 500 mg of flavo-
noids/day confers protection for the endothelial and platelet functions, maintains
blood pressure and blood lipids, and prevents inflammation and thrombosis
(Bondonno et al., 2019). Flavonoids consumption also enhances the cognitive capa-
bilities and restrains the progression of neurodegenerative diseases like Alzheimer’s:
it seems that flavonoids inhibit the neuronal apoptosis induced by inflammation,
reduce the oxidative stress and inhibit key enzymes involved in the fabrication of
amyloid plaques, thus maintaining the number and quality of neurons (Ayaz et al.,
2019; Bakoyiannis et al., 2019). Bioflavonoids are also thought to possess an impor-
tant anticancer activity through the modulation of different enzymes and receptors
involved in cancer development: it seems that flavonoids may inactivate carcino-
gens, reduce proliferation and metastasis, arrest the cell cycle, induce apoptosis,
inhibit angiogenesis, and reverse multidrug resistance process associated with pro-
teins (Ivey et al., 2017; Raffa et al., 2017; Wang et al., 2013). It has been suggested
that flavonoids exert health benefits via antioxidant mechanisms, including direct
scavenging of reactive oxygen species, inhibition of oxidases responsible for super-
oxide anion production, activation of antioxidant enzymes, chelation of metals, and
mitigation of oxidative stress caused by nitric oxide accumulation (Raffa et al.,
2017). Flavonoids exhibited intense antioxidant activity in vitro, but due to their low
bioavailability, the in vivo antioxidant action of flavonoids is indirect and found to
be mediated by cytosolic aryl hydrocarbon receptor (AhR)/nuclear factor erythroid
2-related factor 2 (Nrf2) pathway: flavan-3-ols, flavonols, flavones, and isoflavones
bind to the AhR and induce the translocation of Nrf2 to the nucleus and the tran-
scription of antioxidant enzymes (superoxide dismutase, glutathione peroxidase,
catalase, peroxiredoxin, and heme oxygenase-1) (Pei et al., 2020).
The evidence for flavonoid health benefits accumulated in the last years has
increased the scientific community’s interest in identifying new molecules and
searching for potential extraction sources to develop more effective therapies, new
dietary products, natural food additives, herbal-based pharmaceuticals, and nutra-
ceuticals. To date, interest was focused on the study of flavonoids of green and black
teas (Imran et al., 2018; Raychaudhuri et al., 2019; Stangl et al., 2006), grapes, and
wine (Fernandes et al., 2017; Jenkinson et al., 2012; Tanaka et al., 2019), citrus
fruits (Gandhi et al., 2020; Hwang et al., 2012; Koolaji et al., 2020; Musumeci et al.,
2020), berry fruits (Devore et al., 2012; Esposito et al., 2014; Kalt et al., 2020;
Vauzour et al., 2010), vegetables (Kim et al., 2004; Pérez-Gregorio et al., 2014; Wu
et al., 2019), culinary and medicinal herbs (Coppin et al., 2013; Gutiérrez-Grijalva
et al., 2018; Slimestad et al., 2020; Yoo et al., 2009), cocoa and chocolate (Ahmed
et al., 2020; Martins et al., 2020; Munguia et al., 2020; Socci et al., 2017), but for
the extraction and identification of new molecules, exotic and underutilized fruits,
and vegetables, the wastes and molasses of food and industrialized agricultural
exploitations are considered as interesting alternatives to the food material.
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 421

Flavonoids content in food and plant materials varies widely among the species and
between varieties within the same species. In addition, flavonoid content varies
according to the plant organ, developmental stage, and growth conditions (Petrussa
et al., 2013).

4 Flavonoids in Ficus carica L.

4.1 Total Flavonoids Content in Ficus carica L.

In the data available, flavonoids are found in different parts of F. carica, including
the fruits (peels, pulps, and seeds), leaves, stem bark, and latex in variable amounts.
In 100 g of fresh fruits, total flavonoid content varied considerably between 2.1 mg
(Solomon et al., 2006) to 3694 mg (Mo et al., 2020), but most authors reported val-
ues within the range of 20–900 mg/100 g (Table 19.1). The variability observed in
flavonoid content is related to the fig varieties, the geographical origin of fig sam-
ples, the agro-ecological factors, and the methods used in the extraction, determina-
tion, and expression of results (Aljane et al., 2020; Mahmoudi et al., 2018).
Regarding the dried fig fruits, total flavonoid contents in 100 g of figs vary from
18.07 mg (Bachir Bey et al., 2016) to 275 mg (Soni et al., 2014) (Table 19.1). The
drying process negatively affects the fruit‘s flavonoids, leading to considerable
losses. Sun-drying of figs caused an important decrease in the flavonoid content of
dark fig fruits reaching 86% (Bachir Bey et al., 2016). Sun-drying seems to affect
more purple figs than yellow ones: a loss of 21% of the flavonoid content of purple
figs was reported after drying, while an increase of about 75% was observed in yel-
low figs (Kamiloglu & Capanoglu, 2015). Dark figs are rich in anthocyanins, the
most sensitive class of flavonoids to high temperatures. After drying of dark figs,
anthocyanins disappeared completely, and only traces were detected, while 80%
decrease was reported for flavonols (Bachir Bey et al., 2016).
Fig flavonoids are essentially concentrated in the external parts of the fruit
(peels). Some studies reported the flavonoids content in pulp and peel of fig fruits
separately. For example, 75 mg of flavonoids was found in 100 g of green peels
from Portuguese figs, but only traces of flavonoids were reported in pulps (Palmeira
et al., 2019). When comparing figs with different colors, black figs enclose higher
amounts of flavonoids than green ones. Kamiloglu and Capanoglu (2015) found 234
and 44 mg of flavonoids in 100 g of peels and pulps of purple figs, respectively,
against 63 and 7 mg/100 g of peels and pulps of yellow figs. Given the physiological
role of flavonoids in plants (UV protection of plant cells and nucleic acids) and the
physical protective role of peels for fruit grains, it is logical that flavonoids are con-
centrated in the fruit peels rather than the pulps.
F. carica leaves present higher flavonoid content than fruits (2.62 mg/g of leaves
against 1.96 mg/g of fruits) (Trifunschi et al., 2015). In addition, the amount of
rutin, the most abundant flavonoid compound in F. carica, was found to be two-fold
Table 19.1 Flavonoids extraction from different parts of Ficus carica L
422

Optimal extraction Target

Method Sample procedure Additional information compounds Content References
Conventional methods
SLE Fruit Solvent: MOH 80%, _ TP, TF, AC TP = 69.5 mg GAE/100 g, Aljane et al. (2020)
SSR: 1/25 (g/mL), TF = 7.73 mg QE/100 g,
Time: 2 h (agitation), AC = 4.08 mg CGE/100 g.
Temperature: Ambient.
SLE Fruit Solvent: ETOH 70%, _ TP, TF TP = 10.7 mg GAE/g, Belguith-Hadriche et al.
SSR: 3/100 (g/mL), TF = 5.15 mg CAE/g. (2016)
Time: 24 h (agitation
150 rpm),
Temperature: 37 °C.
SLE Fruit Solvent: ETOH 80%, Optimized parameters: TP, TF, PA TP = 3.7 mg GAE/g, Bucic-Kojic et al. (2011)
SSR: 1/40 (g/mL), – ETOH-H2O ratio, TF = 2.5 mg CE/g,
Time: 120 min (agitation, – temperature. PA = 0.87 mg CyE/g.
200 rpm),
Temperature: 80 °C.
SLE Fruit Solvent: MOH/H2O/HCl _ TP, AC TP = 211.9 mg GAE/100 g, Çalişkan and Polat
(357/17/1.4, v/v/v), AC = 298.9 μg CRE/g. (2011)
SSR: 10/25 (g/mL),
Time: 60 min (agitation),
Temperature: Ambient.
SLE Fruit Solvent: ACN 60%, _ TP, TF TP = 342 mg GAE/100 g, Meziant et al. (2015)
SSR: 1/100 (g/mL), TF = 19.2 mg QE/100 g.
Time: 90 min (agitation),
Temperature: 45 °C.
SLE Fruit Solvent: ACN 60%, _ TP, TF, AC TP = 493.4 mg GAE/100 g, Zidi et al. (2020)
SSR: 1/50 (g/mL), TF = 48.31 mg QE/100 g,
Time: 2 h (agitation), AC = 41.39 mg CGE/100 g.
Temperature: 40 °C.
L. Meziant and M. Bachir-bey
19

SLE Fruit Solvent: ACN 60%, Extraction repeated 2 times. TP, TF, FO, TP = 644 mg GAE/100 g, Bachir bey and
(dried) SSR: 1.34/100 (g/mL), AC, PA TF = 126 mg QE/100 g, Louaileche (2015)
Time: 2 h (agitation FO = 40.42 mg QGE/100 g,
400 rpm), AC = 18.90 mg QGE/100 g,
Temperature: 40 °C. PA = 179.8 mg CyE/100 g.
SLE Fruit Solvent: MOH 80%, _ TP, TF TP = 201.7 mg GAE/100 g, Khadhraoui et al. (2019)
(dried) SSR: 1/6 (g/mL), TF = 112.2 mg QE/100 g.
Time: 60 min (agitation,
200xg),
Temperature: 4 °C.
SLE Fruit Solvent: ACN/DCM/EAC/ _ TP, TF TP = 10.9 mg GAE/g, Soni et al. (2014)
(dried) MOH TF = 2.75 mg CE/g.
(1/1/1/1, v/v/v/v),
SSR: 5/14 (g/mL),
Time: 48 h (agitation,
140 rpm),
Temperature: 40 °C.
SLE Fruit Solvent: ETOH, Optimized parameters: AC AC = 4.03 mg CRE/g. Backes et al. (2018)
peel SSR: 1/20 (g/mL), – extraction time,
Time: 13.74 min (agitation), – temperature,
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Temperature: 35.64 °C. – ETOH-H2O ratio.

SLE Fruit Solvent: MOH 90% + citric Optimized parameters: AC AC = 346.6 mg CGE/100 g. Meziant et al. (2018)
peel acid 5% – solvent,
(10/90, v/v), – SSR,
SSR: 1/100 (g/mL), – time,
Time: 180 min (agitation), – acidification.
Temperature: Ambient. Extraction repeated twice.

(continued)
423
Table 19.1 (continued)
424

Optimal extraction Target

Method Sample procedure Additional information compounds Content References
SLE Fruit Solvent: MOH/H2O/CA 5% Extraction repeated twice. TP, TF, TP = 86.32 mg GAE/g, Meziant et al. (2021)
peel (72/18/10, v/v/v), FO, AC TF = 50.47 mg QE/g,
SSR: 1/100 (g/mL), FO = 37.45 mg RE/g,
Time: 150 min (agitation), AC = 6.22 mg CRE/g.
Temperature: 25 °C.
SLE Fruit Solvent: MOH 50%, Optimized parameters: TP, TF TP = 71.4 mg GAE/100 g, Nakilcioglu-Tas and
seeds SSR: 1/5 (g/mL), – solvent, TF = 31.2 mg CE/100 g. Ôtles (2021)
Time: 90 min (agitation), – solvent/H2O ratio.
Temperature: 50 °C.
SLE Leaf Solvent: H2O or ETOH _ TF Yield = 9% (ETOH). Trifunschi and Ardelean
70%, (2013)
SSR: 1/20 (g/mL),
Time: 60 min (agitation),
Temperature: 60 °C.
SLE Leaf Solvent: H2O, Steam explosion pretreatment: TF TF = 32.73 mg/g Qin and Chen (2015)
SSR: 1/75 (g/mL), – sample rehydration 1/3 (g/ (untreated sample),
Time: 6 h, mL) for 1 h, TF = 51.03 mg/g
Temperature: 80 °C. – steam explosion (0.2– (steam explosion, 0.2 MPa).
0.4 MPa) for 3 min.
Optimized parameters:
– SSR, time, temperature.
– Pretreatment
SLE Leaf + Solvent: MOH 80%, Re-extraction with 150 mL TP, TF, AC TP = 19.7 mg GAE/g (leaf), Boukhalfa et al. (2018)
twig SSR: 15/300 (g/mL), MOH 80% TP = 12.8 mg GAE/g (twig),
Time: 12 h (agitation for 12 h at 40 °C. TF = 9.72 mg QE/g (leaf),
500 rpm), TF = 5.02 mg QE/g (twig),
Temperature: 40 °C. AC = 3.22 mg CGE/100 g
(leaf),
L. Meziant and M. Bachir-bey

AC = 5.12 mg CGE/100 g
(twig).
19

SLE Stem Solvent: H2O, _ TP, TF TP = 133 mg GAE/g, Saoudi and El Feki
SSR: 1/100 (g/mL), TF = 43.2 mg QE/g. (2012)
Time: 15–20 min
(agitation),
Temperature: Ambient.
Maceration Fruit Solvent: ETOH, _ TP, TF, TP = 2190 mg GAE/100 g, Amessis-Ouchemoukh
SSR: 1/50 (g/mL), FO, PA TF = 858 mg QE/100 g, et al. (2017)
Time: 24 h (agitation), FO = 19.5 mg QE/100 g,
Temperature: Ambient. PA = 496.6 mg CE/100 g.
Maceration Fruit Solvent: ETOH 90%, Extraction repeated 4 times. Prenylated Compounds yield: Liu et al. (2019)
SSR: 1/2 (g/mL), Post-extraction fractionation: Isoflavone (1): 0.38%, (2): 0.18%,
Time: 1 week, – extract suspended in H2O Derivatives (3): 0.27%, (4): 1.2%,
Temperature: Ambient. (1/4, w/v), (5): 1.85%, (6): 0.27%,
– partition with PTE (1/1, v/v) (7): 0.13%, (8): 0.18%,
5 times, (9): 2.1%, (10): 1.34%,
– partition with EAC (1/1, (11): 0.26%, (12): 0.58%,
v/v) 5 times. (13): 0.13%, (14): 2.86%,
(15): 0.69%, (16): 0.62%.
Maceration Fruit Solvent: MOH, _ TP, TF TP = 45.8 mg GAE/g, Shahbazi (2017)
SSR: 1/20 (g/mL), TF = 9.88 mg RE/g.
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Time: 12 h,
Temperature: Ambient.
Maceration Fruit Solvent: H2O, ETOH 70%, _ TP, TF, TN TP = 458 mg GAE/100 g Benmaghnia et al. (2019)
(dried) MOH 80% or ACN 50%, (MOH),
SSR: 1/4 (g/mL), TF = 228.2 mg CE/100 g
Time: 24 h (agitation), (ACN),
Temperature: Ambient. TN = 254.1 mg CE/100 g
(H2O).

(continued)
425
Table 19.1 (continued)
426

Optimal extraction Target

Method Sample procedure Additional information compounds Content References
Maceration Fruit Solvent: H2O or PTE, _ TP, TF, TN TP = 756.6 mg GAE/100 g Debib et al. (2014)
(dried) SSR: 1/10 (g/mL), (H2O),
Time: 24 h (agitation), TP = 100 mg GAE/100 g
Temperature: Ambient. (PTE),
TF = 90 mg CE/100 g (H2O),
TF = 58 mg CE/100 g (PTE),
TN = 40 mg GAE/100 g
(ACN),
TN < 10 mg GAE/100 g
(MOH).
Maceration Fruit + Solvent: ETOH 70%, _ TP, TF TP = 25.2 mg CatE/g (leaves), Trifunschi et al. (2015)
Leaf SSR: 2/25 (g/mL), TP = 18.6 mg CatE/g (fruits),
Time: 24 h, TF = 2.62 mg RE/g (leaves),
Temperature: Ambient. TF = 1.96 mg RE/g (fruits).
Maceration Leaf Solvent: MOH, _ TP, TF TP = 96.4 mg GAE/g, Ayoub et al. (2019)
SSR: 1/25 (g/mL), TF = 33.5 mg QE/g.
Time: 48 h (agitation),
Temperature: Ambient.
Maceration Leaf Solvent: H2O or ETOH _ TF Yield = 2.5% (H2O), Trifunschi and Ardelean
70%, Yield = 8% (ETOH). (2013)
SSR: 1/20 (g/mL),
Time: 3 days,
Temperature: Ambient.
Maceration Latex Solvent: MOH, Extraction repeated 3 times, TP, TF TP = 26.8 mg GAE/g, Pasayeva et al. (2020)
SSR: 3/40 (g/mL), Additional solid-phase TF = 8.62 mg CE/g.
Temperature: Ambient. extraction:
Zorbax SPE C18 cartridge.
L. Meziant and M. Bachir-bey
19

Soxhlet Leaf Solvent: MOH, _ TP, TF TP = 49.7 mg GAE/g, Mahmoudi et al. (2016)
extraction SSR: 1/10 (g/mL), TF = 14.1 mg CE/g.
Time: 8 h.
Soxhlet Leaf Solvent: ETOH or MOH or Optimized parameters: TP, TF TP = 150.7 mg GAE/g Radwan et al. (2020)
extraction H2O, – solvent-H2O ratio. (ETOH-butanol fraction),
SSR: 1/25 (g/mL), – fractionation with n-butanol TF = 350 mg RE/g
Time: 2.5 h. (1/1, v/v). (MOH-butanol fraction).
Innovative methods
UAE Fruit Solvent: MOH 80%, _ TP, TF TP = 59 mg GAE/100 g, Marinova et al. (2005)
SSR: 1/100 (g/mL), TF = 20.2 mg CE/100 g.
Time: 20 min,
Ultrasound bath.
UAE Fruit Solvent: H2O, Extraction repeated 3 times. TP, TF TP = 12.5 mg GAE/g, Yang et al. (2009)
SSR: 45/225 (g/mL), Pre-extraction using TF = 1.84 mg CE/g.
Time: 30 mins, supercritical CO2:
Temperature: 25 °C, – lipid extraction (4 h, 45 °C,
Ultrasound bath. 30 MPa,
CO2 flow rate 10 L/h),
– separation (35 °C, 5 MPa).
UAE Fruit Solvent: MOH, Extraction repeated 3 times. TP, TF TP = 181.6 mg/100 g (fresh), Bachir bey et al. (2016)
(fresh SSR: 1/10 (g/mL), TP = 35.75 mg/100 g (dried),
Ficus carica L. as a Source of Natural Bioactive Flavonoids

+ Time: 60 min (first TF = 151.4 mg/100 g (fresh),

Dried) extraction), TF = 18.07 mg/100 g (dried).
30 min (second extraction),
15 min (third extraction),
Ultrasound bath (220 V,
40 kHz).

(continued)
427
Table 19.1 (continued)
428

Optimal extraction Target

Method Sample procedure Additional information compounds Content References
UAE Fruit Solvent: MOH 75% + FA Extraction repeated 3 times. TP, TF, TP = 493 mg GAE/100 g Kamiloglu and
(fresh 0.1% (v/v), PA, AC (fresh), Capanoglu (2015)
+ SSR: 2/5 (g/mL), TP = 417 mg GAE/100 g
dried) Time: 15 min, (dried),
Ultrasound bath. TF = 66 mg CE/100 g (fresh),
TF = 52 mg CE/100 g (dried),
PA = 62 mg CyE/100 g
(fresh),
PA = 16 mg CyE/100 g
(dried),
AC = 83 mg CGE/100 g
(fresh),
AC = 14.5 mg CGE/100 g
(dried).
UAE Fruit Solvent: (MOH 80% or Extraction repeated 3 times. TP, TF, TN TP > 500 mg GAE/100 g Debib et al. (2014)
(dried) ACN 60%) + DTBMP 1%, (ACN),
SSR: 1/5 (g/mL), TP > 400 mg GAE/100 g
Ultrasound bath. (MOH),
TF = 118 mg CE/100 g
(ACN),
TF = 137.4 mg CE/100 g
(MOH),
TN = 100 mg GAE/100 g
(ACN),
TN = 194 mg GAE/100 g
(MOH).
L. Meziant and M. Bachir-bey
19

UAE Fruit Solvent: ETOH, Optimized parameters: AC AC = 4.32 mg CRE/g. Backes et al. (2018)
peel SSR: 1/20 (g/mL), – extraction time,
Time: 21.34 min, – ultrasonic power,
Sonicator (310.58 W). – ETOH-H2O ratio.
IL-UAE Leaf + Solvent: BMIM (1 M), Optimized parameters: Individual Rutin = 222.3 μg/0.1 g Qin et al. (2015)
Fruit SSR: 1/50 (g/mL), – ionic liquid type, phenolic (leaves),
(peel + Time: 30 min, – ionic liquid concentration, compounds, Rutin = 133.7 μg/0.1 g
Pulp) Temperature: 30 °C, – SSR, temperature, including (peels),
Ultrasound bath (50%). – ultrasonic extraction time. Rutin Rutin = 107.9 μg/0.1 g
(pulps).
MAE Fruit Solvent: ETOH, Optimized parameters: AC AC = 4.11 mg CRE/g. Backes et al. (2018)
peel SSR: 1/20 (g/mL), – extraction time,
Time: 5 min, temperature,
Temperature: 62.41 °C, – ETOH-H2O ratio.
Microwave power (400 W).
MAE Leaf Solvent: H2O or ETOH _ TF Yield = 5.5% (H2O), Trifunschi and Ardelean
70%, Yield = 10% (ETOH). (2013)
SSR: 1/20 (g/mL),
Three microwave cycles
3x(3 s power on, 60s power
off),
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Temperature: 70 °C.
Flash Fruit Solvent: ETOH 50%, Optimized parameters: TF TF = 36.94 mg/g Mo et al. (2020).
extraction SSR: 1/50 (g/mL), – solvent concentration,
Time: 80s, – SSR, Extraction time,
Voltage: 150 V. – Extraction voltage.

(continued)
429
Table 19.1 (continued)
430

Optimal extraction Target

Method Sample procedure Additional information compounds Content References
MCAE Fruit Solvent: MOH, _ TP, TF, TP = 74.1 mg GAE/g, Harzallah et al. (2016)
SSR: 1/1 (g/mL), TN, AC TF = 12.7 mg CE/g,
Ultra-Turrax TN = 75.9 mg TAE/g,
homogenization. AC = 33.3 mg CGE/g.
SLE Solid-liquid extraction, UAE Ultrasound-assisted extraction, IL-UAE Ionic liquid-based ultrasound-assisted extraction, MAE Microwave-assisted extrac-
tion, MCAE Mechanochemical-assisted extraction, SSR Solid-to-solvent ratio, MOH Methanol, ETOH Ethanol, ACN Acetone, FA Formic acid, CA Citric acid,
PTE Petroleum ether, DCM Dichloromethane, EAC Ethyl acetate, BMIM1-Butyl-3-methylimidazolium hexafluorophosphate, DTBMP 26-di-tert-butyl-4-
methylphenol, TP Total phenolics, TF Total flavonoids, AC Anthocyanins, FO Flavonols, PA Proanthocyanidins, TN Tannins, GAE Gallic acid equivalents, QE
Quercetin equivalents, CGE Cyanidin-3-glucoside equivalents, CAE Caffeic acid equivalents, CE (+)-catechin equivalents, CyE Cyanidin equivalents, CRE
Cyanidin-3-rutinoside equivalents, QGE Quercetin-3-glucoside equivalents, RE Rutin equivalents, CatE Catechol equivalents, TAE Tannic acid equivalents
L. Meziant and M. Bachir-bey
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 431

higher in leaves than both peels and pulps of fig fruits in the same extraction condi-
tions (Qin et al., 2015). Total flavonoid contents of F. carica leaves reported in the
literature range from 2.62 mg/g (Trifunschi et al., 2015) to 350 mg/g (Radwan et al.,
2020) (Table 19.1). The examination of publications that treated F. carica leaves
revealed many reports about leaf extracts’ pharmacological and antioxidant proper-
ties, but only a few analyzed the flavonoid content. Variety, methods of extraction,
experimental conditions, and expression of results influence considerably the esti-
mation of flavonoid content in F. carica leaves (Konyalιoğlu et al., 2005; Mahmoudi
et al., 2016; Radwan et al., 2020).
Regarding the other parts of F. carica, rare publications were conducted to deter-
mine flavonoid content in fig tree branches. Twig samples from F. carica cultivated
in Algeria, extracted with aqueous methanol (80%), contained 5.02 mg of flavo-
noids per gram of dry matter (Boukhalfa et al., 2018). Aqueous extracts of stem
samples from F. carica grown in Tunisia contain up to 43.2 mg of flavonoids per
gram of dry extract (Saoudi & El Feki, 2012). Flavonoid content of F. carica latex
gathered from the neck of fig fruits was also reported. Fig latex contains 43.2 mg of
total flavonoids per milliliter of the crude sample (Aziz, 2012). In the same line,
latex fractions enclosed between 2.227 mg (n-hexane fraction) and 8.615 mg (solid-
phase extraction) of flavonoids per gram of extract (Paşayeva et al., 2020). To date,
no report has been published on the presence of flavonoids in F. carica roots.

4.2 Flavonoids Extraction from Ficus carica L.

Preparation of bioactive flavonoids from any plant material used as food ingredi-
ents, pharmaceutical, and cosmetic products always begins with an extraction step.
Extraction aims mainly to separate target soluble metabolites from a mixture of
plant compounds, leaving behind the insoluble cellular residue (Azwanida, 2015).
Different extraction methods may be applied, and various strategies depend on the
origin of the biological material from which the target molecules are to be extracted
(Stobiecki & Kachlicki, 2006). Extraction efficiency could be influenced by the
extracting solvent, the solid-to-solvent ratio, time and temperature of extraction,
pH, mode of stirring, number of extractions, and the particle size of plant material;
these parameters influence the recovery of bioactive molecules significantly and
should be considered in optimization procedures (Madureira et al., 2020).
Extraction methods evolved over the years to meet some requirements such as
efficiency, versatility, ease of use, inexpensiveness, and preservation of the major
fraction of bioactive compounds from structural modifications (Gullón et al., 2017).
Conventional methods such as solid-liquid extraction (SLE), maceration, and
Soxhlet extraction are commonly used in bioactive molecules research due to their
simplicity, minimal processing, and the possibility of using several solvents of dif-
ferent polarities. However, they are time-consuming and could cause the degrada-
tion of thermo-sensitive molecules. Also, they are considered non-environmental
respective methods due to large volumes of organic solvents (Azwanida, 2015). To
432 L. Meziant and M. Bachir-bey

overcome the limitations of the conventional methods, innovative extraction meth-

ods are developed, such as ultrasound-assisted extraction (UAE), microwave-
assisted extraction (MAE), enzyme-assisted extraction (EAE), pulsed-electric field
extraction (PEF), mechanochemical-assisted extraction (MCAE), infrared-assisted
extraction (IRAE), pressurized-liquid extraction (PLE), ionic-liquids (ILs), deep
eutectic solvents (DEEs) and supercritical-fluid extractions (SFE). In addition, the
presence of carbohydrates and lipophilic substances in the obtained extracts may
influence the qualitative and quantitative composition of flavonoids and their deriva-
tives, so in many cases, additional cleaning based on solid-phase extraction (SPE)
or liquid-liquid extraction (LLE) of the extracted compounds is required (Stobiecki
& Kachlicki, 2006).
Several extraction procedures extracted flavonoids from F. carica plant material
(Table 19.1). Generally, flavonoids are extracted together with phenolic compounds
(crude phenolic extract), given the similarities in solubility and nature of these com-
pounds. SLE seems to be widely used for the extraction of flavonoids from F. carica
fresh fruits (Aljane et al., 2020; Belguith-Hadriche et al., 2016; Bucic-Kojic et al.,
2011; Çalişkan & Polat, 2011; Meziant et al., 2015; Zidi et al., 2020), dried fruits
(Bachir bey & Louaileche, 2015; Khadhraoui et al., 2019; Soni et al., 2014), fruit
peels (Backes et al., 2018; Meziant et al., 2018; Meziant et al., 2021), fruit seeds
(Nakilcioğlu-Taş & Ötleş, 2021), leaves (Qin & Chen, 2015; Trifunschi & Ardelean,
2013), twigs (Boukhalfa et al., 2018) and stem (Saoudi & El Feki, 2012), using
distilled water, ethanol, methanol, and acetone at different concentrations as extract-
ing solvents with or without heating. Maceration is also used in many extraction
processes for the recovery of flavonoids from F. carica fresh fruits (Amessis-
Ouchemoukh et al., 2017; Liu et al., 2019; Shahbazi, 2017), dried fruits (Benmaghnia
et al., 2019; Debib et al., 2014), leaves (Ayoub et al., 2019; Trifunschi et al., 2015;
Trifunschi & Ardelean, 2013) and latex (Paşayeva et al., 2020). Soxhlet extraction
was only reported for the extraction of flavonoids from F. carica leaves (Mahmoudi
et al., 2016; Radwan et al., 2020). Among the innovative methods, UAE is the most
utilized method for the preparation of flavonoid extracts from F. carica fruits (Bachir
Bey et al., 2016; Backes et al., 2018; Debib et al., 2014; Kamiloglu & Capanoglu,
2015; Marinova et al., 2005; Yang et al., 2009) and leaves (Qin et al., 2015). To a
lesser extent, MAE was also reported to extract flavonoids from F. carica fruit peel
(Backes et al., 2018) and leaves (Trifunschi & Ardelean, 2013). Flavonoids were
also recovered from F. carica fruits by Ultra-Turrax-assisted extraction using meth-
anol as solvent (Harzallah et al., 2016). Recently, a promising and ultra-rapid extrac-
tion method called flash extraction, already used at an industrial scale for the
extraction of pigments from raisin skins for the fabrication of wine, was also assayed
for the extraction of total flavonoids from F. carica fruits using aqueous ethanol
(50%) as extracting solvent, and 80 seconds were sufficient to yield 36.9 mg of
flavonoids from one gram of sample (Mo et al., 2020).
The efficiency of UAE over MAE and SLE for the extraction of cyanidin-3-
rutinoside, the most abundant anthocyanin in dark F. carica fruit peels, was high-
lighted in a comparative study where results showed that UAE produced high purity
extracts with slightly higher anthocyanin content (4.32 mg CRE/g of fig peel)
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 433

compared to the MAE (4.11 mg CRE/g of peel) and SLE (4.03 mg CRE/g of peel)
(Backes et al., 2018). MAE efficiency in extracting flavonoids from F. carica leaves
using aqueous ethanol (70%) was compared to maceration and SLE methods, and
the obtained results showed that MAE provides higher yields (10%, w/w) with the
advantage of saving time (3 cycles of 3 seconds) compared to maceration (8%, w/w)
conducted during 3 days and SLE (9%, w/w) during 60 minutes (Trifunschi &
Ardelean, 2013). The UAE efficiency was compared with the maceration method
for the recovery of phenolic compounds from F. carica leaves, and results showed
that UAE produced higher extraction yields and higher phenolic contents than mac-
eration (10.7–21.1 mg GAE/g for maceration and 12.9–23.0 mg GAE/g for UAE),
but without statistically significant differences (Akhtar et al., 2019). Similarly, UAE
was slightly more effective than maceration for extracting phenolic compounds
from F. carica leaves (0.69 and 0.79 g GAE/100 g for maceration and UAE, respec-
tively) and fruits (0.12 and 0.20 g GAE/100 g for maceration and UAE, respec-
tively). However, both methods gave similar qualitative phenolic profiles (Ammar
et al., 2015). The conventional extraction methods are still widely used in recent
research despite their disadvantages, probably because they provided comparable
extraction efficiency and similar flavonoid profiles as the innovative methods with-
out requiring any specific expensive equipment.

4.3 Flavonoid Profile of Ficus carica L.

4.3.1 Methods of Characterization

After the extraction step necessary to isolate the flavonoid compounds from the plant
material, the study of flavonoids is completed by identification and characterization
steps of the extracted compounds and the elucidation of their structures (aglycones,
glycosylated flavonoids, acylated flavonoids, and isomeric forms). Various analytical
methods could be applied for the identification and quantification purpose, such as
chromatographic systems (ex. thin-layer, liquid, and gas, micellar electro-kinetic,
high-speed counter current, supercritical fluid chromatographies) equipped with dif-
ferent types of analyzers (electromagnetic, Fourier transform ion cyclotron reso-
nance, time of flight, etc.), matrix-assisted laser desorption (MALDI), near-infrared
reflectance spectroscopy (NIRS), capillary zone electrophoresis (CE), etc. (Gullón
et al., 2017). Reverse-phase high performance (RP-HPLC) and ultra-high perfor-
mance (UHPLC) liquid chromatographies coupled to UV-Visible spectrophotometry
and mass spectrometry (MS) are established as the methods of choice for the separa-
tion and quantification of several individual flavonoids simultaneously by compari-
son of retention times as well as spectral properties to purified known standards. The
characterization and identification of unknown compounds are performed with the
aid of nuclear magnetic resonance (NMR) analyses (1H and 13C isotopes) which are
required to provide structural information (Stobiecki & Kachlicki, 2006).
Research studies about the flavonoid profile of extracts of different F. carica
parts and their main findings are assembled in Table 19.2. Most research works were
Table 19.2 Flavonoid compounds identified in Ficus carica L. extracts
434

Sample Extract Characterization Identified compounds Quantification References

Fruits (fresh) Acetone HPLC-UV Quercetin 0.655 mg/100 g Gündesli et al. (2021)
25% + 1% and -DAD Kaempferol 2.396 mg/100 g
trifluoroacetic Myricetin 2.632 mg/100 g
acid
Naringenin 0.054 mg/100 g
Catechin 0.590 mg/100 g
Epicatechin 7.809 mg/100 g
Fruits (fresh) Methanol UHPLC- Quercetin 9.48–145.5 mg/kg Soltana et al. (2018)
QTOF-MS Quercetin 3-O-rutinoside (rutin) 107.7–303.2 mg/kg
Quercetin 3-O-glucoside 5.20–69.1 mg/kg
(isoquercetin)
Quercetin 3-O-(6″-malonyl) 0.16–1.90 mg/kg
glucoside
Quercetin di-glycoside derivatives 2.74–20.7 mg/kg
Kaempferol 1.39–21.5 mg/kg
Kaempferol 3-O-glucoside 1.10–15.34 mg/kg
Isorhamnetin 3-O-glucoside 0.03–0.24 mg/kg
Syringetin 0.03–0.09 mg/kg
Tamarixetin 0.01–0.13 mg/kg
Luteolin 8-C-glucoside (orientin) 6.80–38.6 mg/kg
Luteolin 7-O-glucoside (cynaroside) 2.37–15.1 mg/kg
Luteolin C-glycoside derivatives 5.9–32.9 mg/kg
Apigenin 8-C-glucoside (vitexin) 14.53–71.1 mg/kg
Apigenin C-glycoside derivatives 6.04–20.6 mg/kg
Eriodictyol 7.07–25.5 mg/kg
Naringenin 7.13–44.4 mg/kg
L. Meziant and M. Bachir-bey
19

Fruits (fresh) Ethanol 70% RP-UHPLC-DAD Quercetin 2.0% (relative area). Belguith-Hadriche et al.
-QTOF-MS/MS Quercetin 3-O-rutinoside (rutin) 53.9%. (2016)
Quercetin 3-O-glucoside 2.5%.
(isoquercetin)
Quercetin 0.4%.
3-O-(6″-malonyl)-glucoside
Apigenin 8-C-glucoside (vitexin) 5.0%.
Apigenin C-glycoside derivatives 3.5%.
Luteolin 0.9%.
Luteolin 8-C-glucoside (orientin) 1.2%.
Luteolin 6-C-glucoside (isoorientin) 0.8%.
Naringenin 0.5%.
Eriodictyol 0.2%.
Taxifolin 1.4%.
Cyanidin 3,5-diglucoside 8.4%.
Cyanidin 3-rutinoside 17.6%.
Fruits (fresh) Methanol 30% + UHPLC-PDA Kaempferol 3-O-rutinoside 0.6–30.5 mg/100 g Wojdylo et al. (2016)
1% ascorbic acid -QTOF/MS Quercetin 3-O-rutinoside 39.0–328 mg/100 g
Quercetin 3-O-glucoside 4.6–14.2 mg/100 g
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Quercetin 3-O-(malonyl) glucoside 5.3–58.0 mg/100 g

Apigenin C-glycoside 0.5–24.3 mg/100 g
Cyanidin 3-O-rutinoside 0.4–116 mg/100 g
Cyanidin 3,5-O-diglucoside 0.0–1.9 mg/100 g
Pelargonidin 3-O-rutinoside 0.0–3.8 mg/100 g
(+)-Catechin 20.6–95.7 mg/100 g
(−)-Epicatechin 9.0–43.1 mg/100 g
Procyanidins 55.6–268 mg/100 g
435

(continued)
Table 19.2 (continued)
436

Sample Extract Characterization Identified compounds Quantification References

Fruits (fresh) Methanol + HPLC-DAD Rutine 4.89–28.7 mg/100 g Veberic et al. (2008)
1% BHT (+)-Catechin 1.07–4.03 mg/100 g
(−)-Epicatechin 0.34–0.97 mg/100 g
Fruits (syrup) Methanol + HPLC-DAD-MS Rutin 1.9 mg/g Puoci et al. (2011)
1% BHT Catechin 0.34 mg/g
Epicatechin 0.007 mg/g
Fruit seeds Methanol 50% HPLC-DAD Rutin 50.7 mg/kg DM Nakilcioglu-Tas & Ötles
(−) Epicatechin 166.4 mg/kg DM (2021)
Fruits (dried) Methanol 80% LC-ESI-MS Quercetin 0.24–1.49 mg/100 g Khadhraoui et al. (2019)
Quercetin 3-O-rutinoside (rutin) 2.17–38.05 mg/100 g
Kaempferol 0.02–1.13 mg/100 g
Apigenin 0.00–4.85 mg/100 g
Apigenin 7-O-glucoside 0.00–0.91 mg/100 g
Luteolin 0.00–0.15 mg/100 g
Luteolin 7-O-glucoside 0.02–0.43 mg/100 g
Cirsiliol 0.95–2.72 mg/100 g
Naringenin 0.01–0.43 mg/100 g
Naringenin 7-rhamnoglucoside 0.00–0.06 mg/100 g
(naringin)
Fruits (dried) Ethyl acetate HPLC and 1H and Ficucaricone D (Not reported). Liu et al. (2019)
13
fraction from C NMR Gancaonin N (Not reported).
ethanol 90% Isopiscerythrone (Not reported).
extract
Viridiflorin (Not reported).
4’-Hydroxy-5,7-dimethoxy-6-(3-methyl- (Not reported).
2-butenyl)-isoflavone
L. Meziant and M. Bachir-bey
Fruits (dried) Methanol 20% RP-HPLC-DAD Rutin 3.26 mg/100 g DM Tawfik and Alhejy (2014)
19

Quercetin 0.56 mg/100 g DM

Kaempferol 0.12 mg/100 g DM
Isorhamnetin 0.29 mg/100 g DM
Catechin 13.8 mg/100 g DM
Epicatechin 12.8 mg/100 g DM
Epigallocatechin 25.4 mg/100 g DM
Epicatechin gallate 0.85 mg/100 g DM
Epigallocatechin gallate 2.52 mg/100 g DM
Fruits (dried) Aqueous HCl HPLC-DAD Quercetin 3-O-rutinoside 4.33–88.40 mg/kg Faleh et al. (2012)
(pH = 2) Quercetin 0.10–8.76 mg/kg
Fruits (fresh Methanol HPLC-DAD Quercetin 6.76–16.5 mg/100 g (fresh), Bachir bey et al. (2016)
+ dried) 0.64–3.43 mg/100 g (dried).
Quercetin 3-rutinoside (rutin) 31.7–38.1 mg/100 g (fresh),
3.49–10.8 mg/100 g (dried).
Quercetin 3-glucoside (isoquercetin) 5.03–14.1 mg/100 g (fresh),
0.49–3.20 mg/100 g (dried).
Cyanidin 3-glucoside 1.34–6.86 mg/100 g (fresh).
Cyanidin 3-rutinoside 31.4–73.0 mg/100 g (fresh),
0.00–0.57 mg/100 g (dried).
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Catechin 1.72–8.63 mg/100 g (fresh),

1.80–5.78 mg/100 g (dried).
Procyanidin B1 1.87–4.80 mg/100 g (fresh),
0.77–1.56 mg/100 g (dried).
Procyanidin B2 1.11–2.61 mg/100 g (fresh),
0.47–1.34 mg/100 g (dried).
Fruits (fresh Methanol + HPLC-DAD Rutin 15.1–68.2 mg/100 g (fresh), Nakilcioglu & Hisil
+ dried) 1% BHT 5.75–12.35 mg/100 g (dried). (2013)
(−) Epicatechin 42.2–76.9 mg/100 g (fresh),
14.7–23.4 mg/100 g (dried).
437

(continued)
Table 19.2 (continued)
438

Sample Extract Characterization Identified compounds Quantification References

Fruits (fresh Methanol + HPLC-DAD Rutin 0.61–1.86 mg/100 g (fresh), Slatnar et al. (2011)
+ dried) 1% BHT + and -ESI-MS/MS 1.38–12.06 mg/100 g (sun dried),
3% formic acid 3.75–14.62 mg/100 g (oven dried).
Quercetin 3-O-glucoside 0.18–0.60 mg/100 g (fresh),
0.56–3.35 mg/100 g (sun dried),
1.10–2.98 mg/100 g (oven dried).
Kaempferol 3-O-glucoside 0.04–0.13 mg/100 g (fresh),
0.31–0.59 mg/100 g (sun dried),
0.56–1.43 mg/100 g (oven dried).
Luteolin 8-C-glucoside 0.00 mg/100 g (fresh),
0.13–0.16 mg/100 g (sun dried),
0.21–0.45 mg/100 g (oven dried).
Cyanidin 3-O-rutinoside 0.21–0.62 mg/100 g (fresh),
0.12–0.26 mg/100 g (sun dried),
0.12–0.31 mg/100 g (oven dried).
Catechin 1.36–2.88 mg/100 g (fresh),
5.88–11.4 mg/100 g (sun dried),
15.5–19.7 mg/100 g (oven dried).
Epicatechin 7.11–8.67 mg/100 g (fresh),
10.4–23.3 mg/100 g (sun dried),
26.6–36.6 mg/100 g (oven dried).
Fruits (peel Ethanol 80% LC-DAD-ESI-MS/ Quercetin 3-O-rutinoside (rutin) 0.513 mg/g ext. (peel). Palmeira et al. (2019)
+ pulp) MS Quercetin acetylglycoside-derivatives 0.038 mg/g ext. (peel).
Kaempferol glycoside derivatives Traces (pulp).
Apigenin 0.026 mg/g ext. (peel).
2”-O-rhamnose-C-acetylglycoside
Apigenin C-glycosylated derivatives 0.072 mg/g ext. (peel).
Taxifolin glycoside derivative 0.097 mg/g ext. (peel).
L. Meziant and M. Bachir-bey
19

Fruits (peel Methanol 80% HPLC-DAD Quercetin 3-rutinoside (rutin) 14.2–15.3 mg/100 g (peel), Viuda-Martos et al. (2015)
+ pulp) 0.08–0.10 mg/100 g (pulp).
Luteolin 1.07–2.82 mg/100 g (peel),
0.05–0.06 mg/100 g (pulp).
Luteolin 7-O-glucoside 2.18–3.80 mg/100 g (peel).
Apigenin 1.03–5.10 mg/100 g (peel),
0.10–0.11 mg/100 g (pulp).
Cyanidin 3-rutinoside 0.43–2.01 mg/100 g (peel).
Cyanidin 3-glucoside 0.12–1.33 mg/100 g (peel).
Catechin 6.10–12.48 mg/100 g (peel),
4.58–9.53 mg/100 g (pulp).
Epicatechin 2.25–5.74 mg/100 g (peel),
1.68–1.75 mg/100 g (pulp).
Fruits (peel Methanol HPLC-DAD Quercetin 3-O-rutinoside (rutin) 527.9–1071 mg/kg (peel). Del Caro and Piga (2008)
+ pulp) Flavonol compound I 54.82–83.52 mg/kg (peel).
Flavonol compound II 114.6–296.1 mg/kg (peel).
Cyanidin 3-O-rutinoside 901.0 mg/kg (peel),
4.92–17.6 mg/kg (pulp).
Cyanidin 3-O-glucoside 28.24 mg/kg (peel).
Ficus carica L. as a Source of Natural Bioactive Flavonoids

(continued)
439
Table 19.2 (continued)
440

Sample Extract Characterization Identified compounds Quantification References

Fruits (peel Methanol +0.5% HPLC-DAD-MS Cyanidin 3-rutinoside 1.542–7.832 mg/100 g (peel), Duenas et al. (2008)
+ pulp) Trifluoroacetic 0.104–1.016 mg/100 g (pulp).
acid Cyanidin 3-rutinoside dimer 0.044–0.093 mg/100 g (peel),
0.00–0.008 mg/100 g (pulp).
Cyanidin 3-glucoside 0.151–1.538 mg/100 g (peel),
0.019–0.215 mg/100 g (pulp).
Cyanidin 3-malonylglucoside 0.067–0.351 mg/100 g (peel),
0.002–0.009 mg/100 g (pulp).
Cyanidin 3, 5-diglucoside 0.199–0.523 mg/100 g (peel),
0.009–0.057 mg/100 g (pulp).
Cyanidin 0.053–0.139 mg/100 g (peel),
3-malonylglycosyl-5-glucoside 0.002–0.009 mg/100 g (pulp).
5-Carboxypyrano-Cyanidin 0.048–0.126 mg/100 g (peel),
3-rutinoside 0.004–0.094 mg/100 g (pulp).
Pelargonidin 3-glucoside 0.029–0.068 mg/100 g (peel),
0.001–0.009 mg/100 g (pulp).
Pelargonidin 3-rutinoside 0.047–0.352 mg/100 g (peel),
0.002–0.008 mg/100 g (pulp).
(Epi)catechin-(4 → 8)-Cyanidin 0.00–0.048 mg/100 g (peel),
3-glucoside 0.003–0.013 mg/100 g (pulp).
(Epi)catechin-(4 → 8)-Cyanidin 0.059–0.123 mg/100 g (peel),
3-rutinoside I 0.002–0.059 mg/100 g (pulp).
(Epi)catechin-(4 → 8)-Cyanidin 0.044–0.093 mg/100 g (peel),
3-rutinoside II 0.002–0.013 mg/100 g (pulp).
(Epi)catechin-(4 → 8)-Pelargonidin 0.00–0.024 mg/100 g (peel).
3-rutinoside I
(Epi)catechin-(4 → 8)-Pelargonidin 0.00–0.020 mg/100 g (peel).
3-rutinoside II
L. Meziant and M. Bachir-bey
19

Fruits (peel Acetone/water HPLC-DAD- Quercetin rutinoside 2.9–15.8 mg/100 g (peel), Vallejo et al. (2012)
+ pulp + /formic acid ESI-MS 0.0–1.8 mg/100 g (pulp),
dried) (80/19.5/0.5) and -MS/MS. 10.2–13.0 mg/100 g (dried).
Quercetin glucoside 0.2–3.2 mg/100 g (peel),
0.7–2.5 mg/100 g (dried).
Quercetin acetylglucoside 0.0–1.7 mg/100 g (peel),
2.1–2.8 mg/100 g (dried).
Kaempferol rutinoside 0.2–0.9 mg/100 g (peel),
1.0–2.0 mg/100 g (dried).
Apigenin 7-O-rutinoside 0.8–2.5 mg/100 g (peel).
Luteolin C-glycosides derivatives 0.1–1.9 mg/100 g (peel).
Cyanidin 3-rutinoside 0.0–108.9 mg/100 g (peel),
0.0–9.5 mg/100 g (pulp).
Cyanidin 3-glucoside 0.0–11.1 mg/100 g (peel).
Proanthocyanidins 0.1–1.0 mg/100 g (pulp).
Catechin/epicatechin 9.0–38.7 mg/100 g (pulp).
Fruits (fresh) Ethanol 70% HPLC-DAD Quercetin 0.08% (fruits), 2.5% (leaves). Trifunschi et al. (2015)
+ Leaves Rutin 1.8% (fruits), 0.2% (leaves).
Kaempferol Traces (fruits + leaves).
Ficus carica L. as a Source of Natural Bioactive Flavonoids

Luteolin Traces (fruits), 0.07% (leaves).

Fruits (dried) Aqueous HPLC-DAD Quercetin 0.14 mg/100 g (leaves). El-Shobaki et al. (2010)
+ leaves Galangin 24.76 mg/100 g (fruits).
Chrysin 7.64 mg/100 g (fruits),
7.14 mg/100 g (leaves).
Pinocembrin 46.94 mg/100 g (fruits),
52.85 mg/100 g (leaves).
Pinostrobin 19.47 mg/100 g (leaves).

(continued)
441
Table 19.2 (continued)
442

Sample Extract Characterization Identified compounds Quantification References

Fruits (peel Methanol 80% RP-UHPLC-DAD- Quercetin (relative abundance): Ammar et al. (2015)
+ pulp) + Ethanol 70% QTOF 0.69–1.06% (fruits),
Leaves -MS and -MS/MS 0.59–1.34% (peel),
0.33–0.86% (pulp).
Quercetin 3-O-rutinoside (rutin) 16.34–19.20% (fruits),
31.41–43.68% (peel),
0.71–0.75% (pulp),
14.46–14.96% (leaves).
Quercetin 3-O-glucoside 1.95–2.04% (fruits),
(isoquercetin) 2.55–3.17% (peel),
0.86–1.05% (pulp),
4.28–4.67% (leaves).
Quercetin 3-O-(6‘’-malonyl) 0.48–1.60% (fruits),
glucoside 1.45–3.00% (peel),
0.24–0.30% (pulp),
0.49–1.54% (leaves).
Quercetin O-glycoside derivatives 1.88–3.57% (fruits),
1.93–3.51% (peel),
0.50–0.55% (pulp),
0.72–1.27% (leaves).
Luteolin 0.75–0.76% (fruits),
0.50–1.07% (peel),
1.51–2.35% (pulp),
0.37–0.56% (leaves).
Luteolin 8-C-glucoside (orientin) 0.71–0.91% (fruits),
1.20–1.35% (peel),
0.49–0.55% (pulp),
0.92–1.59% (leaves).
L. Meziant and M. Bachir-bey
19

Luteolin 6-C-glucoside (isoorientin) 0.40–0.87% (fruits),

0.97–1.01% (peel),
0.35–0.85% (pulp),
0.45–0.68% (leaves).
Luteolin 7-O-glucoside (cynaroside) 0.19–0.37% (fruits),
0.36% (peel),
0.62–0.72% (pulp),
0.17–0.24% (leaves).
Luteolin C-glycoside derivatives 0.50–0.53% (fruits),
0.52–0.63% (peel),
2.46–3.14% (leaves).
Apigenin 0.90–1.94% (fruits),
0.74–0.89% (peel),
1.32–1.48% (pulp),
0.14–0.18% (leaves).
Apigenin 8-C-glucoside (vitexin) 2.06–3.29% (fruits),
2.57–3.69% (peel),
1.26–1.91% (pulp),
1.53–4.21% (leaves).
Apigenin C-glycoside derivatives 2.66–8.71% (fruits),
4.53–9.68% (peel),
Ficus carica L. as a Source of Natural Bioactive Flavonoids

1.06–1.37% (pulp),
18.7–19.1% (leaves).
Eriodictyol glycoside derivatives 0.71–2.30% (fruits),
2.03–5.12% (peel),
0.58–0.87% (pulp).
Eriodictyol 0.73–1.46% (fruits),
0.81–3.41% (peel),
0.38–0.70% (pulp).

(continued)
443
Table 19.2 (continued)
444

Sample Extract Characterization Identified compounds Quantification References

Naringenin 0.71–1.60% (fruits),
0.78–1.72% (peel),
0.61–0.79% (pulp),
0.06–0.07% (leaves).
Taxifolin (dihydroquercetin) 0.42–0.52% (fruits),
0.50–0.66% (peel),
0.79–0.95% (pulp).
Genistein 0.09–0.14% (leaves).
Genistein 4′-methyl ether (biochanin 0.09–0.74% (fruits),
A) 0.08–0.14% (peel),
0.08–0.24% (pulp),
0.25–0.50% (leaves).
Hydroxygenistein 0.01–0.03% (leaves)
methyl-ether-malonylglycoside
7-Methoxy 2′-hydroxy genistein 0.73–1.40% (fruits),
(cajanin) 0.19–0.20% (peel),
1.37–4.97% (pulp),
0.49–1.09% (leaves).
Prenylgenistein I / II/ III 2.67–6.27% (fruits),
0.40–1.17% (peel),
7.56–17.89% (pulp),
4.57–6.92% (leaves).
Prenylhydroxygenistein I / II / III 3.79–12.1% (fruits),
0.65–0.71% (peel),
29.8–30.9% (pulp),
8.24–10.6% (leaves).
Cyanidin rutinoside I / II / III 1.85–52.7% (fruits),
33.2–82.5% (peel),
5.02–22.5% (pulp).
L. Meziant and M. Bachir-bey
19

Cyanidin 3-rutinoside-glycoside 1.07% (fruits),

2.25% (peel).
Cyanidin 3,5-diglucoside 1.87–9.71% (fruits),
5.97–19.6% (peel),
5.19–24.7% (pulp).
Cyanidin 1.04% (fruits),
3-malonylglucosyl-5-glucoside 0.83% (peel),
1.34% (pulp).
Cyanidin 3-glucoside 1.70% (fruits),
1.70% (peel),
2.13% (pulp).
Petunidin + dimer cyanidin rutinoside 2.49% (fruits),
3.01% (peel).
Pelargonidin 3-rutinoside 2.00% (fruits),
2.83% (peel).
(+) Catechin 0.17–0.20% (fruits),
0.06–0.07% (peel),
0.27–0.72% (pulp),
0.03–0.04% (leaves).
Fruits (peel BMIM PF6 HPLC-UV Rutin 1.34 mg/g (peel), Qin et al. (2014)
Ficus carica L. as a Source of Natural Bioactive Flavonoids

+ pulp) + 1.08 mg/g (pulp),

Leaves 2.22 mg/g (leaves).
Fruits (peel Aqueous HPLC-DAD Quercetin 3-rutinoside 629.6 mg/kg ext. (peel), Oliveira et al. (2009)
+ pulp) + 64.6 mg/kg ext. (pulp),
Leaves 17,440 mg/kg ext. (leaves).
Quercetin 3-glucoside 30.8 mg/kg ext. (peel),
611 mg/kg ext. (leaves).

(continued)
445
Table 19.2 (continued)
446

Sample Extract Characterization Identified compounds Quantification References

Leaves Ethyl acetate GC-HRMS Quercetin, Kaempferol, Chrysin (Not reported). Dureshahwar et al. (2019)
Fraction of
Methanolic
extract
Leaves Ethanol 70% RP-UHPLC-MS Quercetin 3-O-rutinoside (rutin) 19.8–23.0% (relative area). Belguith-Hadriche et al.
and -MS/MS Quercetin 3-O-glucoside 2.1–2.9%. (2017)
(isoquercetin)
Apigenin 8-C-glucoside (vitexin) 1.0–1.2%.
Apigenin C-glycoside derivatives 8.1–12.7%.
Luteolin 0.8–2.5%.
Luteolin C-glycoside derivatives 1.8–2.4%.
7-Methoxy 2′-hydroxy genistein 1.2–1.4%.
(cajanin)
Prenylgenistein 10.2–11.9%.
Prenylhydroxygenistein 8.7–10.2%.
Leaves Water/methanol/ HPLC-DAD-MS/ Quercetin 3-O-rutinoside (rutin) 4.7–14.6 mg/g Takahashi et al. (2014)
Acetone (1/1/1) MS Quercetin 3-O-glucoside (Not reported). and (2017)
(isoquercetin)
Quercetin (Not reported).
3-O-(6”-O-malonyl)-glucoside
Kaempferol 3-O-glucoside (Not reported).
(astragalin)
Isoschaftoside 2.5–6.4 mg/g
Leaves Methanol 70% HPLC-DAD and Quercetin 3-O-rutinoside (rutin) 1233 mg/100 g Teixeira et al. (2006,
LC-ESI-MS/MS 2009)
L. Meziant and M. Bachir-bey
19

Leaves Ethanol 70% HPLC-DAD and Quercetin 630 mg/kg extract. Vaya and Mahmood
LC-ESI-MS Luteolin 680 mg/kg extract. (2006)
Biochanin A 17.4 mg/kg extract.
BHT 2,6-di-tert-butyl 4-methyl phenol, BMIM PF6 1-butyl-3-methylimidazolium hexafluorophosphate, GC-HRMS, gas chromatography coupled to high resolution
mass spectrometry, LC-ESI-MS liquid chromatography coupled to electrospray ionization mass spectrometry, RP-HPLC-DAD reversed phase high performance liquid
chromatography with diode array detection, HPLC-UV high performance liquid chromatography with ultraviolet detection, HPLC-DAD-ESI-MS/MS high perfor-
mance liquid chromatography with diode array detection coupled to electrospray ionization and tandem mass spectrometry, RP-UHPLC-DAD-QTOF-MS/MS reversed
phase ultra high performance liquid chromatography with diode array detection coupled to quadrupole time-of-flight tandem mass spectrometry, UHPLC-PDA-
QTOF-MS ultrahigh performance liquid chromatography with photodiode array detection coupled to quadrupole time-of-flight mass spectrometry
Ficus carica L. as a Source of Natural Bioactive Flavonoids
447
448 L. Meziant and M. Bachir-bey

carried out on fruit extracts (fresh, dried, peels and pulps, juices and seeds) and
leaves to a lesser extent, but no research was found on the identification of flavo-
noids in other plant parts (branches, wood, stem bark, roots, and latex). The analyti-
cal methods applied to Ficus carica extracts are mainly HPLC and UHPLC methods
with diode array detectors (Bachir Bey et al., 2016; Del Caro & Piga, 2008;
El-Shobaki et al., 2010; Faleh et al., 2012; Gündeşli et al., 2021; Nakilcioğlu &
Hışıl, 2013; Nakilcioğlu-Taş & Ötleş, 2021; Oliveira et al., 2009; Tawfik & Alhejy,
2014; Trifunschi et al., 2015; Veberic et al., 2008; Viuda-Martos et al., 2015) or UV
detector (Gündeşli et al., 2021; Qin et al., 2015). Higher sensitivity, more compound
specificity, and larger flavonoid profiles are achieved by coupling the HPLC or
UHPLC to mass spectrometry (Dueñas et al., 2008; Puoci et al., 2011), a tandem
mass spectrometry (Belguith-Hadriche et al., 2017; Slatnar et al., 2011; Takahashi
et al., 2014, 2017; Vallejo et al., 2012), an electrospray ionization mass spectrome-
try (Khadhraoui et al., 2019; Palmeira et al., 2019; Slatnar et al., 2011; Teixeira
et al., 2006, 2009; Vallejo et al., 2012; Vaya & Mahmood, 2006) and a quadrupole
time-of-flight mass spectrometry (Ammar et al., 2015; Belguith-Hadriche et al.,
2016; Soltana et al., 2018; Wojdyło et al., 2016). Flavonoids from ethyl acetate
extract of Ficus carica leaves were separated for the first time using gas chromatog-
raphy equipped with high-resolution mass spectrometry (Dureshahwar et al., 2019).
The major drawback of this method is that only identification of flavonoid agly-
cones is possible, and the intact flavonoid conjugates did not appear in the profile
obtained since these compounds undergo derivatization before GC analysis.

4.3.2 General Flavonoid Profile

Six flavonoid subclasses were reported in F. carica extracts, namely flavonols, fla-
vones, isoflavones, flavanones, flavan-3-ols, and anthocyanins. The qualitative and
quantitative variations in flavonoid profiles reported in the literature are liable to the
cultivars used, the seasonal differences, and agricultural land type and practices
(Gündeşli et al., 2021). The flavonoid profile of F. carica also depends on the plant
part considered in the study. Flavones are predominant in leaves with relative abun-
dance between 26.1% (green varieties) and 28.35% (dark varieties), followed by
flavonols (20.5–21.8%) and isoflavones (13.6–19.3%) (Ammar et al., 2015).
Besides, flavonols are the major flavonoids in fruits of green varieties with a relative
abundance of 21.3%, followed by flavones (17.3%), while anthocyanins predomi-
nate in fruits of dark varieties with a relative abundance of 70.8%, followed by fla-
vonols (27.4%) and isoflavones (19.8%) (Ammar et al., 2015). Differences in
flavonoid profiles are also reported between pulps and peels within the same variety.
In effect, the repartition of flavonoids between these two parts is characterized by a
high abundance of anthocyanins (52.8–99.1%), flavonols (38.6–53.9%), and fla-
vones (17.3–18.3%) in peels, and a high abundance of isoflavones (42.6–50.2%) in
pulps (Ammar et al., 2015). Recently, in singular research, flavonoids belonging to
the subclasses of flavonols and flavan-3-ols were reported to occur also in fig seeds,
with flavan-3-ols content being three-times higher than flavonols (Nakilcioğlu-Taş
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 449

& Ötleş, 2021). The drying process of figs was reported to affect different sub-
classes of flavonoids. After sun drying of figs, flavan-3-ols decreased from
9.87 mg/100 g (mean value) to 5.53 mg/100 g, and flavonols decreased from
53.7 mg/100 g (mean content of 3 dark varieties) to 11.2 mg/100 g. In comparison,
anthocyanins showed a severe degradation from 58.75 mg/100 g (mean value) to
less than 0.57 mg/100 g or disappeared completely after the sun-drying process,
indicating the high thermo-sensitivity anthocyanins (Bachir Bey et al., 2016).

4.3.3 Flavonols in Ficus carica L.

Flavonols (3-hydroxy-2-phenylchromen-4-one) constitute the major group of flavo-

noid compounds distributed extensively in nature. They are characterized by a C2-C3
double bond and a hydroxyl group at C3 with an Oxo group at C4. The multiple
hydroxyl, methoxyl, and glycoside groups on the basic structure give differences in
biological and pharmacological properties of flavonols (Sharma et al., 2018). These
compounds are found in their aglycone form and are also frequently glycosylated
by attaching a monosaccharide (ex. glucose, rhamnose) or a disaccharide (ex. rutin-
ose) to the hydroxyl group in position 3 of the C ring. The chemistry of flavonols
provides them with high antioxidant activity; in addition to other astonishing prop-
erties such as the protective effect on the cardiovascular system, inhibition of key
enzymes (adenosine deaminase, tyrosine kinase, aldose reductase, etc.), regulation
of some hormones (estrogen, androgen, thyroid hormones, etc.), reduction of
inflammation, decrease in blood cell aggregation, inhibition of microbial growth,
and other activities (Brahmachari & Gorai, 2006).
Flavonols occur massively in fruits of F. carica as quercetin and kaempferol
derivatives. Quercetin aglycone was identified in fresh fruits (Ammar et al., 2015;
Bachir Bey et al., 2016; Belguith-Hadriche et al., 2016; Gündeşli et al., 2021;
Soltana et al., 2018; Trifunschi et al., 2015), dried fruits (Bachir Bey et al., 2016;
Faleh et al., 2012; Khadhraoui et al., 2019; Tawfik & Alhejy, 2014) and leaves
(Ammar et al., 2015; Dureshahwar et al., 2019; El-Shobaki et al., 2010; Trifunschi
et al., 2015; Vaya & Mahmood, 2006). Among quercetin derivatives, quercetin-3-O-
rutinoside (rutin) is the predominant compound found in considerable amounts in
F. carica fruit peels and leaves, with relative abundance up to 43.68% in peels and
14.96% in leaves (Ammar et al., 2015). Rutin is known to possess an intense anti-
oxidant activity against free radicals and lipid peroxidation (Yang et al., 2008),
important antimicrobial (Al-Majmaie et al., 2019), and anti-inflammatory and neu-
roprotective activities (Hao et al., 2016). Rutin was reported to have anticancer
activity through different mechanisms, including cell growth and proliferation inhi-
bition, apoptosis and autophagy induction, angiogenesis inhibition, carcinogen
inactivation, and cytochrome P450 modulation, in addition to a high chemopreven-
tive potential (Farha et al., 2020). Rutin is currently marketed as an active com-
pound in many dietary supplements and prescribed to treat varicose veins, internal
bleeding, hemorrhoids, etc. Oral doses of rutin (from 500 mg to 2000 mg/day) are
taken for extended periods (up to 6 months) without any notable side effects (Gullón
450 L. Meziant and M. Bachir-bey

et al., 2017). Other quercetin derivatives were also identified in F. carica extracts
(Table 19.2), namely quercetin-3-O-glucoside (isoquercetin) (Ammar et al., 2015;
Bachir Bey et al., 2016; Belguith-Hadriche et al., 2016, 2017; Oliveira et al., 2009;
Slatnar et al., 2011; Soltana et al., 2018; Takahashi et al., 2014;Vallejo et al., 2012;
Wojdyło et al., 2016), quercetin-3-O-(6″-malonyl) glucoside (Ammar et al., 2015;
Belguith-Hadriche et al., 2016; Soltana et al., 2018; Takahashi et al., 2014; Wojdyło
et al., 2016), quercetin O-glycosides (Ammar et al., 2015), quercetin di-glycosides
(Soltana et al., 2018), and quercetin acetyl-glycosides (Palmeira et al., 2019; Vallejo
et al., 2012). Kaempferol aglycone was identified in F. carica fresh fruits (Gündeşli
et al., 2021; Soltana et al., 2018; Trifunschi et al., 2015), dried fruits (Khadhraoui
et al., 2019; Tawfik & Alhejy, 2014), and leaves (Dureshahwar et al., 2019;
Trifunschi et al., 2015). Two kaempferol glycosides were identified in F. carica
extracts, namely kaempferol 3-O-glucoside (astragalin) (Slatnar et al., 2011; Soltana
et al., 2018; Takahashi et al., 2014) and kaempferol 3-O-rutinoside (Vallejo et al.,
2012; Wojdyło et al., 2016). Myricetin was reported recently for the first time in
fresh figs and revealed to be the most abundant flavonol aglycone (2.632 mg/100 g)
(Gündeşli et al., 2021). Galangin (El-Shobaki et al., 2010) and isorhamnetin (Tawfik
& Alhejy, 2014) were only identified in dried figs. In addition to the compounds
mentioned above, minor flavonols, namely syringetin, tamarixetin and
isohamnetin-3-O-glucoside were also reported in the methanolic extract of fresh
fruits (Soltana et al., 2018).

4.3.4 Flavones in Ficus carica L.

Flavones (2-phenylchromen-4-one) are the principal pigments in white and cream-

colored flowers and act as co-pigments with anthocyanins in blue flowers (Hostetler
et al., 2017). Flavones and flavonols share the same structure and chemical proper-
ties except for the absence of the hydroxyl group at C3. Flavones occur widely in
nature as 7-O-glycosides and 6-C- and 8-C-glycosides, where the glycoside moiety
is directly attached to the aromatic carbon atom, the thing that is not found in flavo-
nols (Hollman & Arts, 2000). Natural flavones show several biological activities,
including antioxidant, DNA protective, cardiovascular protective, anticancer, antip-
roliferative, anti-inflammatory, antimicrobial, anti-HIV, and anti-histaminic activi-
ties (Brahmachari & Gorai, 2006).
The main flavones found in F. carica are apigenin and luteolin in aglycone and
glycosylated forms. Glycosylated compounds found in F. carica plant parts pre-
sented mostly C glycosidic links at positions 6, 7, and 8 with glucose, rhamnose,
arabinose, or rutinose-O liked glycosides (exclusively with glucose) are also found
at position 7. Apigenin derivatives are more abundant in fruits and leaves than luteo-
lin derivatives (Table 19.2). Apigenin aglycone was found in fruits in abundance
(0.90–1.94%) (Ammar et al., 2015); it is found in peels in higher amounts compared
to pulps (Viuda-Martos et al., 2015). Apigenin glycosides were found in abundance
in leaves: apigenin 8-C-glucoside (vitexin) (1.53–4.21%), apigenin C-diglycoside
derivatives (18.7–19.1%) (Ammar et al., 2015) and apigenin-6-C-
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 451

arabinoside-8-C-glucoside (isochaftoside) (2.5–6.4 mg/g) (Takahashi et al., 2014).

Apigenin-7-O- glucoside was reported in dried figs (Khadhraoui et al., 2019).
Apigenin-7-O-rutinoside was identified in fig peels (Vallejo et al., 2012). Traces of
an acetyl-glycosylated apigenin compound (apigenin-(2”-O-rhamnoside)-C-acetyl
glycoside) was also reported in fig peels (Palmeira et al., 2019). Regarding luteolin,
the aglycone was found to occur in F. carica fresh fruits (Ammar et al., 2015;
Belguith-Hadriche et al., 2016; Viuda-Martos et al., 2015), dried fruits (Khadhraoui
et al., 2019), and leaves (Ammar et al., 2015; Belguith-Hadriche et al., 2017;
Trifunschi et al., 2015; Vaya & Mahmood, 2006). Luteolin glycosides identified in
F. carica extracts are (1) luteolin-7-O-glucoside (cynaroside) found in fresh fruits
(Ammar et al., 2015; Soltana et al., 2018; Viuda-Martos et al., 2015), in dried fruits
(Khadhraoui et al., 2019), and leaves (Ammar et al., 2015), (2) luteolin-8-C-gluco-
side (orientin) found in fresh fruits (Ammar et al., 2015; Belguith-Hadriche et al.,
2016; Soltana et al., 2018), in dried fruits (Slatnar et al., 2011), and leaves (Ammar
et al., 2015), and (3) luteolin-6-C-glucoside (isoorientin) found in fresh fruits
(Ammar et al., 2015; Belguith-Hadriche et al., 2016) and leaves (Ammar et al.,
2015). Other C-glycosylated luteolin derivatives (structures are not fully elucidated
yet) were reported in fresh figs (Ammar et al., 2015; Soltana et al., 2018; Vallejo
et al., 2012) and leaves (Ammar et al., 2015; Belguith-Hadriche et al., 2017). Along
with apigenin and luteolin, two other flavones were found in F. carica, chrysin was
identified in dried fruits (El-Shobaki et al., 2010), and leaves (Dureshahwar et al.,
2019; El-Shobaki et al., 2010), and cirsiliol was reported to occur in dried figs
(Khadhraoui et al., 2019).

4.3.5 Isoflavones in Ficus carica L.

Isoflavones (3-phenylchromen-4-one) are a critical biologically active subclass of

flavonoids that occur primarily in the Fabaceae family, particularly soybeans, con-
sidered the principal source of dietary isoflavones. These compounds showed poten-
tial antioxidant, estrogenic, LDL-cholesterol reducing, cardio-vascular protective,
and anticancer activities (Vitale et al., 2013).
Isoflavones are reported in F. carica fruits and leaves, primarily as genistein and
2′-hydroxygenistein derivatives (aglycone and prenylated forms) (Table 19.2).
Genistein aglycone and a glycosylated derivative of hydroxygenistein (hydroxygen-
istein methyl ether-malonylglycoside) were found exclusively in leaves (Ammar
et al., 2015), while 4′-methyl ether genistein (biochanin A) and 7-methoxy-2′-hyd
roxygenistein (cajanin) were found in fig fruits (in both peels and pulps) (Ammar
et al., 2015) and leaves (Ammar et al., 2015; Belguith-Hadriche et al., 2017; Vaya
& Mahmood, 2006). Different isomeric forms of prenylgenistein and prenylhydrox-
ygenistein were reported to occur in fig fruits (more abundant in pulps than in peels)
(Ammar et al., 2015) and also in leaves (Belguith-Hadriche et al., 2017). Besides,
other prenylated isoflavones were described for the first time in F. carica fruits, i.e.,
gancaonin N, isopiscerythrone, viridiflorin, 4′-hydroxy-5,7-dimethoxy-6-(3-meth
452 L. Meziant and M. Bachir-bey

yl-2-butenyl) isoflavone, in addition to a newly reported isoflavone named ficucari-

cone D, isolated from ethyl acetate extract fractionated on silica gel column chro-
matography and identified using the 1H and 13C NMR (Liu et al., 2019).

4.3.6 Anthocyanins in Ficus carica L.

Anthocyanins (2-phenyl-chromen-1-ylium) are the most colorful group of flavo-

noids. These pigments, contained in the flowers and the external part of fruit peels,
confer the red, blue, purple, and black coloring depending on pH: in acidic condi-
tions, anthocyanins appear as red pigments, while in alkaline conditions, anthocya-
nins show blue pigments. Anthocyanins are present in the reddish fruit pulps and
leaves, grains, and tubers. Nevertheless, these pigments are highly sensitive to tem-
perature, light, and pH (Khoo et al., 2017). Besides being strong antioxidants,
anthocyanins and anthocyanidins (the aglycone forms of anthocyanins) are poten-
tial pharmaceutical ingredients, possessing various medicinal properties, including
antidiabetic, anticancer, anti-inflammatory, anti-obesity, antimicrobial, and cardio-
protective activities (Khoo et al., 2017).
Anthocyanins are found exclusively in F. carica fruits, distinctively abundant in
peels of dark varieties. Most anthocyanins described in figs are cyanidin and pelar-
gonidin derivatives (Table 19.2). Cyanidin-3-O-rutinoside is considered the main
anthocyanin compound in figs, with content varying between 0.21 and
116.2 mg/100 g in fresh fruits (Ammar et al., 2015; Bachir Bey et al., 2016;
Belguith-Hadriche et al., 2016; Slatnar et al., 2011; Solomon et al., 2006; Wojdyło
et al., 2016), between 0.12 and 0.57 mg/100 g in dried fruits (Bachir Bey et al.,
2016; Slatnar et al., 2011), between 0.43 and 108.9 mg/100 g in peels (Ammar
et al., 2015; Del Caro & Piga, 2008; Dueñas et al., 2008; Vallejo et al., 2012; Viuda-
Martos et al., 2015), and between 0.10 and 9.5 mg/100 g in pulps (Ammar et al.,
2015; Del Caro & Piga, 2008; Dueñas et al., 2008; Vallejo et al., 2012). Dimers of
cyanidin 3-rutinoside and other derivatives (cyanidin 3-rutinoside-glycoside and
5-carboxypyrano-cyanidin-3-rutinoside) were also reported in figs (Ammar et al.,
2015; Dueñas et al., 2008). In addition, cyanidin-3,5-O-diglucoside and cyanidin-3-
O-glucoside are found to be more abundant in fig pulps (5.19–24.74% and 2.13%,
respectively) than in peels (5.97–19.6% and 1.70%, respectively) (Ammar et al.,
2015). Other cyanidin derivatives were identified in fig peels and pulps, including
cyanidin-3-malonylglucoside (Dueñas et al., 2008) and cyanidin-3-
malonylglucosyl-5-glucoside (Ammar et al., 2015; Dueñas et al., 2008). Pelargonidin
derivatives described in fig peels and pulps are pelargonidin-3-O-rutinoside (Dueñas
et al., 2008) and pelargonidin-3-O-glucoside (Ammar et al., 2015; Dueñas et al.,
2008; Wojdyło et al., 2016). The only anthocyanidin found in figs was petunidin,
identified as a dimer of cyanidin rutinoside and reported for the first time in the
peels of a dark fig variety (Ammar et al., 2015).
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 453

4.3.7 Flavanones in Ficus carica L.

Another subclass of flavonoids present in F. carica plant parts is flavanones. These

compounds are the Citrus flavonoids, given their high prevalence in Citrus species
(oranges, limes, lemon, mandarins, tangors, grapefruits, etc.) in their extracts and
juices. Compounds belonging to the flavanones group share the basic structure
(2-phenylchroman-4-one) that lacks the double bond in position C2-C3. This speci-
ficity in structure gives a non-planar molecular orientation to flavanones, and this
feature could impact their biological properties and their interaction with biological
receptors compared to flavonols and flavones. Different biological activities were
reported for flavanone compounds, including antioxidant, anti-inflammatory, anti-
cancer, anti-proliferative, anti-cholesterolemic, antimicrobial, and cardio-protective
activities (Barreca et al., 2017; Khan et al., 2014).
The main flavanones described in F. carica plant parts are naringenin, taxifolin,
and eriodictyol derivatives (Table 19.2). Naringenin aglycone was reported in fresh
fruits (Ammar et al., 2015; Belguith-Hadriche et al., 2016; Gündeşli et al., 2021;
Soltana et al., 2018), dried fruits (Khadhraoui et al., 2019), and leaves (Ammar
et al., 2015). The only naringenin glycoside found in dried figs is naringin
(naringenin-7-rhamnoglucoside) (Khadhraoui et al., 2019). Taxifolin (dihydroquer-
cetin) aglycone was reported in fresh figs (in both peels and pulps) (Ammar et al.,
2015; Belguith-Hadriche et al., 2016), while taxifolin glycoside derivatives were
reported only in peels (Palmeira et al., 2019). Regarding eriodictyol, the aglycone
was identified in fresh figs (Ammar et al., 2015; Belguith-Hadriche et al., 2016;
Soltana et al., 2018), and glycoside derivatives were reported in fig peels and pulps
(Ammar et al., 2015). In a single study, two additional flavanones were identified in
F. carica leaves, namely pinostrobin (19.47 mg/100 g) and pinocembrin
(52.8 mg/100 g); the latter also occurs in fruits (46.94 mg/100 g) (El-Shobaki
et al., 2010).

4.3.8 Flavan-3-Ols in Ficus carica L.

The last flavonoid subgroup found in F. carica extracts is the group of flavan-3-ols
(2-phenyl-3,4-dihydrochromen-3-ol) that constitute 83.5% of the total dietary flavo-
noids and are found in abundance in tea, cocoa, grapes, and apples (Lei et al., 2016).
Flavan-3-ols form a diversified group of compounds with different molecular
weights, from simple monomers such as catechin and epicatechin to complex poly-
mers known as proanthocyanidins (Aron & Kennedy, 2008). Several biological
properties were attributed to flavan-3-ols, such as antioxidant, anti-inflammatory,
anti-cancer, antidiabetic, antibacterial, immuno-modulatory, and cardiovascular
protective (Aron & Kennedy, 2008; Lei et al., 2016; Márquez Campos et al., 2020;
Mena et al., 2014; Nawrot-Hadzik et al., 2021; Raman et al., 2018).
All flavan-3-ols identified in F. carica are catechin and epicatechin derivatives
(Table 19.2). (+)-Catechin and (−)-epicatechin monomers were reported in fresh
figs (Bachir Bey et al., 2016; Gündeşli et al., 2021; Nakilcioğlu & Hışıl, 2013; Puoci
454 L. Meziant and M. Bachir-bey

et al., 2011; Slatnar et al., 2011; Veberic et al., 2008; Viuda-Martos et al., 2015;
Wojdyło et al., 2016) and dried figs (Bachir Bey et al., 2016; Nakilcioğlu & Hışıl,
2013; Slatnar et al., 2011; Tawfik & Alhejy, 2014; Vallejo et al., 2012). In fruit
seeds, only (−)-epicatechin was reported in a higher amount (16.64 mg/100 g) than
rutin (5.07 mg/100 g) (Nakilcioğlu-Taş & Ötleş, 2021). A small amount of (+)-cat-
echin (0.04%) was also found in F. carica leaves (Ammar et al., 2015). Dimers of
catechin and epicatechin formed with anthocyanins (i.e., cyanidin-3-glucoside,
cyanidin-3-rutinoside, and pelargonidin-3-rutinoside) were also described in fig
peels and pulps (Dueñas et al., 2008). Epicatechin gallate, epigallocatechin, and
epigallocatechin gallate were reported in dried figs 7-times higher than flavonols
(Tawfik & Alhejy, 2014). Regarding polymeric flavan-3-ols, up to 268 mg/100 g of
procyanidins were reported in fresh figs (Bachir Bey et al., 2016; Vallejo et al.,
2012; Wojdyło et al., 2016) and dried figs (Bachir Bey et al., 2016).

5 Bioaccessibility of Flavonoids in Ficus carica L.

The availability of flavonoids after digestion is crucial as the reduced bioavailability

of certain antioxidants is considered a restraining aspect of their potential health
benefits. The bioavailability of flavonoids depends on multiple factors. The first fac-
tor is the macro and micro-nutrients that surround flavonoids in the gastrointestinal
tract: sugar increases considerably the bioavailability of flavan-3-ols in chocolate
matrix, vitamin C increases the intestinal absorption of catechins in green tea, vita-
min E enhances the bioavailability of anthocyanins in fruits and lipids increases
both intestinal micellar absorption of flavonoids and plasma concentration of flavo-
noid metabolites; on the other hand, dietary fibers reduce the intestinal absorption
of flavonoids and proteins interact with flavonoids and form complexes that decrease
their bioavailability. The second factor is the glycosylation of flavonoids: flavonoid
aglycones are lipophilic and could pass the intestinal epithelial cells via passive dif-
fusion, while glycosylated flavonoids need epithelial transporters that limit their
intestinal absorption. The third factor is the metabolic transformations: the forma-
tion of methyl, sulfate, and glucuronic derivatives of flavonoids after absorption in
the liver and kidney could influence their biological activities. Finally, the last factor
is the type of flavonoids: the highest bioavailability is recorded for quercetin gluco-
sides, catechin, flavanones, and isoflavones, while anthocyanins and proanthocyani-
dins present the lowest bioavailability (Kamiloglu et al., 2020; Kopustinskiene
et al., 2020).
Flavonoids‘bioaccessibility differs from one species to another. For example, in
the case of Ficus carica, recent research investigated the in vitro oro-gastrointestinal
digestibility of 3 varieties of fresh and dried figs and their effect on total flavonoids
and proanthocyanidin contents (Kehal et al., 2021). Results showed that the diges-
tive process (oral, gastric, and intestinal phases) caused a gradual decrease in total
flavonoid content of the fig varieties from 54–98% (before digestion) to 18–37%
(after the intestinal phase) in fresh figs, and from 58–98% (before digestion) to
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 455

27–32% (after the intestinal phase) in dried figs, on the contrary, proanthocyanidins,
in both fresh and dried figs, increased of 12–35% after the oral phase, then decreased
gradually in the gastric and intestinal phases. Similarly, antioxidant activity was
affected by the gastrointestinal digestion; fresh figs lost 69–82% of their initial
reducing power and 39–54% of their radical scavenging activity, while losses esti-
mated to 77–85% and 49–55% of the initial reducing power and radical scavenging
activity, respectively were found in dried figs at the end of the intestinal phase of
digestion. Whatever the nature of the figs, the decrease in flavonoid content during
the digestive process was explained by an enzymatic degradation by PPO and per-
oxidases present in the samples and by the instability of these compounds in the
simulated digestive conditions (mechanical action, enzymatic activity, and pH
effect) that led to hydrolysis of the most sensitive compounds, in addition to the fact
that gastrointestinal digestion release other compounds present in figs (ex. proteins
and sugars) which interact with flavonoids and cause changes in structure and solu-
bility leading to poor extractability.
Earlier, in vitro gastrointestinal digestion on antioxidant and some phenolic com-
pounds of fresh and dried figs was determined (Kamiloglu & Capanoglu, 2013).
Results showed that the initial antioxidant activity decreased considerably after gas-
trointestinal digestion by 74% and 83% in yellow and purple whole figs. Among the
investigated compounds, rutin, cyanidin-3-glucoside (C3G), and cyanidin-3-
rutinoside (C3R) were tested, and results revealed that only 12% and 16% of the
initial rutin content was recovered after intestinal digestion of whole fresh yellow
and purple figs, respectively. Although anthocyanin compounds (C3G and C3R)
undergo a partial gastric degradation (39%) in yellow figs, an increase in anthocy-
anin contents was observed for dried purple figs, explained by the effect of pH and
pepsin digestion that increase the flavylium cation in the solution. However, antho-
cyanins were barely detectable or not observed at all in the dialyzed intestinal frac-
tions of purple figs (losses attained 95–100%), a situation explained by the
transformation of the red flavylium cation formed in the gastric phase at pH = 1.7 to
the colorless chalcone at pH = 7 in the intestinal phase, or by oxidation or degrada-
tion into other chemicals which cannot be detected.
The in vitro gastrointestinal digestibility is helpful as a rapid, safe, and ethically
acceptable tool to study the bioaccessibility of food elements, but the generated
results may not reflect the real in vivo digestion conditions, given the complexity of
biological interactions, the physiological absorptions in the stomach and small
intestine, and the transformations by the microbiota in the colon, conditions that are
not taken into consideration in the in vitro simulated experiments. There is an appar-
ent controversial situation between anthocyanins‘real health beneficial effects and
their low bioavailability (generally measured with in vitro digestibility assays). It
was found that intact anthocyanins are the major form detected in blood; such a situ-
ation was explained by a specific mechanism of absorption or metabolism suggest-
ing that anthocyanins may be absorbed directly from the stomach by transport
through bilitranslocase at the gastric level because anthocyanins are good ligands
for this carrier (Manach et al., 2004).
456 L. Meziant and M. Bachir-bey

6 Impact of Post-harvest Processing on Flavonoids in Ficus

carica L.

Fresh figs are highly perishable fruits because of their soft and delicate skin tissue
(easily damaged), their climacteric behavior with moderate ethylene respiration
(fruits continue to ripen and produce ethylene after harvest), and also because of
their high humidity and nutrient contents (sugars, amino acids, minerals, etc.) which
encourages microbial growth and spoilage, as a consequence, the shelf-life of fresh
figs is limited. As a result, the commercial-quality is reduced (Zidi et al., 2020). In
order to extend the shelf-life of fig fruits and avoid spoilage, the producers generally
apply post-harvest drying of fruits as a practical and economical method of preser-
vation: humidity of fresh fruits is reduced to below 26% by sun-drying, oven-drying,
micro-wave drying, freeze-drying or osmotic dehydration (Bachir Bey et al., 2016;
Naikwadi et al., 2010; Sharifian et al., 2012; Xanthopoulos et al., 2010). However,
some of these methods lead to physic-chemical modifications of the fruits (non-
enzymatic browning, color modification, etc.) and degradation of some thermal-
sensitive bioactive compounds. Therefore, alternative methods of preserving fig
fruits were proposed, such as edible coatings, modified atmosphere packaging, and
product transformations (fig marmalade, juice, wine, etc.) to preserve the visual
aspect of fresh figs and avoid chemical modifications and bioactive compounds
degradation.
The effect of modified atmosphere packaging (MAP) on fresh fig bioactive com-
pounds and antioxidant activity was investigated by Zidi et al. (2020). Flavonoid
content decreased significantly in figs conditioned under MAP by about 38% after
10 days of storage; while a significant increase in anthocyanins content (23.65%)
was reported for figs stored for 5 days under MAP as a result of continued matura-
tion and respiration processes of the fruits at first, followed by a drop of 39% of
anthocyanins content after 10 days of storage explained by the instability of these
compounds under high CO2 and low O2 concentrations. As a consequence of flavo-
noid decrease, the free radical scavenging activity and chelating capacity of figs
stored under MAP were decreased by about 76% and 18%, respectively. The authors
concluded that besides the extended shelf-life of figs for up to 10 days and freshness
maintained, the MAP technique significantly limited the ripening process by retard-
ing the degradation of bioactive compounds and maintaining antioxidant activity
during the storage time (10 days).
In an attempt to extend the shelf-life of fresh figs and maintain the nutritional
quality and the antioxidant function, Aloe vera gel at different concentrations (25,
50, and 75%) was applied as a coating of fresh figs cold-stored for 15 days
(Mirshekari et al., 2020). Aloe vera gel coating at 50% and 75% caused a significant
increase in anthocyanins content (from 11.12 to 21.45 mg Cyanidin-3-O-glucoside
Eq./100 g FW) of coated fresh figs after 15 days of storage; in addition to a higher
total flavonoid content and antioxidant activity. These observations were explained
by the persistence of flavonoids and anthocyanins biosynthesis in fig fruits after
harvesting and that the Aloe vera gel coating activated enzymes involved in the
19 Ficus carica L. as a Source of Natural Bioactive Flavonoids 457

biosynthesis process like phenylalanine ammonia-lyase (PAL). Results proved that

the application of Aloe vera gel as a coating treatment to fresh figs could be a valu-
able alternative to the currently used preserving protocols in maintaining the fresh-
ness of figs for 15 days and preserving at the same time flavonoids antioxidant
capacity of figs.

7 Concluding Remarks

The present chapter reviewed numerous scientific reports searching for the flavo-
noids found in Ficus carica plant parts. As a result, more than 70 compounds
belonging to six different subclasses of flavonoids were identified in the fruits and
leaves of F. carica, with appreciable contents that could form a solid base for indus-
trial exploitation. Food industries that transform fig fruits into ready-to-use products
(jam, syrup, confectioneries, wine & liquors, etc.) generate considerable quantities
of co-products, including the peels (that contain high amounts of flavonols and
anthocyanins), the whole non-conform and overripe fruits discarded during the pro-
cessing that could be exploited as a valuable material for the flavonoids extraction.
Moreover, a great variety of different figs currently exist, and each of them pos-
sesses its flavonoid profiles and, therefore, their adequate uses. Besides, exploiting
the flavonoids of fig leaves could be more beneficial in pharmaceuticals. In contrast,
fig fruits, which require further handing out (compound isolation, purification, and
encapsulation to protect the compounds from degradation and improve their bioac-
cessibility), could be the base of high value-added nutraceutical production.
Nonetheless, there is a need to fill the flagrant lack in flavonoids research observed
in the literature published to date on the F. carica plant parts (latex, root, branches,
stem, wood, etc.), and this for a complete vision of the real potential of an astonish-
ing plant widely spread and used as food and medicine.

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Chapter 20
Fig Minerals

Sarfaraz Ahmed Mahesar, Hadia Shoaib, Abdul Rauf Khaskheli,

Syed Tufail Hussain Sherazi, Abdul Hameed Kori, and Niaz Ali Malghani

1 Introduction

Ficus carica (F. carica) is a small deciduous tree native to Asia Minor, Persia, Syria,
and the Mediterranean region known as Fig (Abu-Mustafa et al., 1963). Ficus
belongs to the Moraceae family, which includes many hemiepiphytes, climbers, and
creepers native to the tropical and subtropical regions. Figs are grown in several
countries worldwide, including Italy, Turkey, Egypt, Morocco, Spain, Greece, the
United States, and Brazil (Frodin, 2004; Rahmani & Aldebasi, 2017). Turkey is still
the world's leading fig producer and exporter, with an average of 269,126 tonnes per
year from 2007 to 2017 and exporting nearly 75% of its production (Turco
et al., 2020).
Figs are commonly consumed both fresh and dried. Drying fruits for later use is
common because they have a limited post-harvest shelf life of 7–8 days if refriger-
ated (Veberic et al., 2008). In terms of nutritional content, dried figs are the best of
the dried fruits, providing a good supply of fiber, minerals, and vitamins ( Badgujar
et al., 2014). In addition, proteins, amino acids, polyphenols, carbohydrates, and
lipids are all present in this food, low in fat and cholesterol. The fig cultivars and
genotypes have a significant impact on the concentrations of these chemicals
(Solomon et al., 2006; Veberic & Mikulic-Petkovsek, 2016). Fresh and dried figs
and fig jam or chocolate coating are common ingredients in Mediterranean desserts

S. A. Mahesar (*) · H. Shoaib · S. T. H. Sherazi · A. H. Kori · N. A. Malghani

National Centre of Excellence in Analytical Chemistry, University of Sindh,
Jamshoro, Sindh, Pakistan
e-mail: [email protected]
A. R. Khaskheli
Institute of Pharmacy, Shaheed Mohtarma Benazir Bhutto Medical University,
Larkana, Sindh, Pakistan

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 467
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_20
468 S. A. Mahesar et al.

such as Panettone and Colombia, often served around Christmas and Easter (Turco
et al., 2020). It is an essential part of the Mediterranean diet because of the high
concentrations of nutrients, vitamins, minerals, amino acids, and antioxidant sub-
stances like phenolic acids it contains (Amessis-Ouchemoukh et al., 2017; Bucic-
Kojic et al., 2011; Oliveira et al., 2010).
Fig tree products have numerous therapeutic properties, such as the fruit's anti-
angiogenic, anticancer, and anti-inflammation properties, which are well-known in
traditional medicine. These include the treatment of cardiovascular, respiratory, and
gastrointestinal disorders, including colitis, indigestion, nausea, and diarrhoea
(Rubnov et al., 2001), antibacterial activity (Aref et al., 2010), cytotoxicity activity
(Khodarahmi et al., 2011), antihelmintic activity (Amol et al., 2010), antimutagenic
activity (Agabeĭli & Kasimova, 2005), antipyretic activity (Patil et al., 2010), anti-
spasmodic activity, antiplatelet activity (Gilani et al., 2008), antiviral activity
(Lazreg Aref et al., 2011), immunostimulant activity (Patil et al., 2010), anti-
inflammatory and antispasmodic remedy (Amessis-Ouchemoukh et al., 2017; Idrus
et al., 2018; Mawa et al., 2013), hypoglycemic activity (El-Shobaki et al., 2010),
and hypolipidemic activity (Asadi et al., 2006).
Individuals who prefer a vegan diet or who have medical conditions that prevent
them from consuming dairy or meat products, such as a milk allergy, may use fruits
like pomegranates and olives as an alternative osteoporosis nutritional supplement
(Chin & Ima-Nirwana, 2016; Nazrun Shuid & Naina Mohamed, 2013). It has been
stated that the mineral composition of figs is similar to human milk, with iron as the
essential mineral. The iron level is also believed to be 50% higher than beef liver
(Lydia, 2009).
Several religious texts, including the Bible and the Quran, reference the herb’s
medicinal benefits. For example, in the Quranic sura “I swear by the fig and the
olive,” these plants’ medical benefits are mentioned (Sura no. 95 verse no. 1).
Furthermore, the Prophet Mohammad (Peace Be Upon Him) advised eating figs to
treat piles and rheumatism, according to Hazrat Abu Darda R.H. (Farooqi, 1998).

2 Botanical Description

A big deciduous tree or large shrub (15–30 ft) with numerous branches from a sin-
gle trunk is the most common form of F. carica. The tree’s bark can be grey, smooth,
dull-white, or hairy. Oval or nearly orbicular, the blades are 10–20 cm long, the peti-
ole is 5–7.5 cm long, and the leaves are rough above and hairy below. Unisexual,
tiny flowers cluster on the inner surface of a wide, hollow, outwardly pear-shaped
receptacle. The receptacle is held at the base by several broad, smooth, spiky bracts,
and the apex is perforated by an opening closed by numerous thin scales.
Male flowers are scarce and can be found near the bottom of the receptacles.
Stamens: 1–5, usually three; sepals: four; filaments: longer than perianth. Female
flower: Pedicellate, sepals-4, perianth very delicate and transparent, deeply cut into
20 Fig Minerals 469

Fig. 20.1 The ripe and unripe fig fruit. (Frodin, 2004; Rahmani & Aldebasi, 2017).

3–5 acute segments, ovary superior, hyaline, 1-celled with a single ovule, found on
the upper part of receptacles, short-stalked.
To distinguish it from other ficus species, the fruit of F. carica has a pear-shaped
interior cavity that produces an array of tiny fruits, which are similar in shape and
size to the syconium of those other species (Salma et al., 2020). Figure 20.1 shows
the ripe and unripe fruit of fig.
There are four basic pomological classifications of figs, depending on the nature
of the flowers and the mode of pollination, Common fig, Capri fig, Smyrna fig, and
San Pedro fig. The Smyrna fig is the most commercially important of the four vari-
eties, and it is widely grown in the USA and Europe (Al-Snafi, 2017; Badgujar
et al., 2014; Salma et al., 2020)
Detailed botanical information is discussed below (Joseph & Raj, 2011).
Kingdom: Plantae
Division: Magnoliophyta
Class: Magnoliopsida
Order: Rosales
Family: Moraceae
Genus: Ficus
Species: Ficus carica

3 Ingredients of F. carica

Fruits and leaves of F. carica contain a wide range of beneficial compounds.

Phenolic compounds, organic acids, and volatile chemicals are found in most F. car-
ica species (Gibernau et al., 1997; Oliveira et al., 2009). Insoluble carbohydrates,
protein, vitamin A, vitamin C, calcium (Ca), and iron (Fe) (Teixeira, Patão, Coelho,
& da Costa) can all be found in fruits, including cyanidin-3-O-glucoside, cyanidin-3-
Orhamnoglucoside, saturated fat, cholesterol, and sodium (Na) (Oliveira et al.,
2009; Teixeira et al., 2006; Solomon et al., 2006). Researchers found that dried fruit
470 S. A. Mahesar et al.

Table 20.1 Chief ingredients of F. carica

Parts of
F. carica Constituent References
Fruits Cyanidin-3-O-glucoside, cyanidin-3-Orhamnoglucoside, Teixeira et al. (2006)
saturated fat, cholesterol, Na, insoluble sugars, protein,
vitamin A, vitamin C, Ca, Fe
Ripe Alkaloids, flavonoids, coumarins, saponins, and terpenes Abu-Mustafa et al.
dried (1963) and Vaya and
fruit Mahmood (2006)
Dried Fixed oil, including fatty acids Oliveira et al. (2009)
seeds
Leaves Furanocoumarins including psoralen, bergapten, xanthotoxin, Ahmed et al. (1990),
triterpenes such as calotropenyl acetate and lupeol acetate. It Çalişkan and Polat
also includes 3-O- and 5-O-caffeoylquinic acids, ferulic acid, (2011), and Oliveira
quercetin-3-O-glucoside, quercetin-3-O-rutinoside and et al. (2009)
psoralen

extracts contain alkaloids, flavonoids, coumarins, and saponins(Teixeira et al.,

2006; Vaya & Mahmood, 2006). Furanocoumarins such as psoralen, bergapten,
xanthotoxin, and triterpenes such as calotropenyl acetate and lupeol acetate have
been found in the leaves (Abu-Mustafa et al., 1963; Ahmed et al., 1990). Phenolics,
anthocyanins, fructose, glucose, and sucrose were also found in figs and a prior
study found fruit to contain phytosterols (Çalişkan & Polat, 2011; Jeong & Lachance,
2001) (Table 20.1).

4 Mineral Composition of Fig

Minerals are inorganic substances that play an essential role in human and animals'
health and disease states and are necessary for almost all metabolic functions in liv-
ing cells (Ndukwe et al., 2013). Macro and micro minerals can be classified based
on their natural distribution. For example, Na, Potassium (K), Ca, and Magnesium
(Mg) are macro-minerals, while Chromium (Cr), Copper (Cu), Fe, Manganese
(Mn), and Zinc (Zn) are micro-minerals. The minerals can be synthesized as conju-
gated proteins and/or attached to other micro molecules, i.e., phosphates, phytates,
polyphenols, etc. (Sulaiman et al., 2011). Figure 20.2 shows the classification of
macro- and micro-elements.
Fe is required to transport oxygen, oxidative metabolism, and cell development
(Özcan & Akbulut, 2008). It is mostly absorbed in the duodenum, carried through
the circulation and extracellular fluid bound to transferrin, and stored intracellularly
in the form of ferritin in humans (Nnorom et al., 2007). Likewise, Cu and Cr are
essential in various physiological processes (Karadaş & Kara, 2012; Khan et al.,
2014). Zn is essential for cellular reproduction and the development of the immune
system (Başgel & Erdemoğlu, 2006; Salgueiro et al., 2002).
20 Fig Minerals 471

Fig. 20.2 Classification of minerals (macro- and micro-elements)

Many chronic, epidemic, endemic, and even malignant disorders are linked to
specific elements, such as Cu, Fe, and Zn, which assist fight infection (Bogden &
Oleske, 2007; Heghedus-Mindru et al., 2014). Anemia, for example, affects one-
third of the world's population due to a deficit in Fe. Enzymes require trace elements
to function properly. Therefore, a trace element deficit will result in a specific condi-
tion that reflects the nutrient's specific functions in the animal’s metabolism
(Mahmood et al., 2012; Duran et al., 2008).
These nutrients are generally received through diet, although not enough is
obtained in most cases. Prelatent, latent, and manifested metal inadequacies are
caused by poor dietetic habits and unhealthy food intake. There are many commer-
cial supplements on the market today, but few of them contain the necessary ingre-
dients in the appropriate natural form; thus, their absorption or bioavailability in the
human body is not as intended (Ficsor et al., 2013).
Figs are the richest source of fiber, vitamin K, and minerals such as Cu, Mn, Mg,
K, Ca, Fe, Na, Phosphorus (P), and Cr adequate for people’s needs, as per USDA
mission variety data. Because 5 dkg (dekagram) of dried figs provides more than
15% of a person's daily need for nutrients. The elements listed below are found in
many sections of the fig, including the leaves, stems, and fruits, Mg, Mn, Fe, Ca, Cu,
Na, K, P, Selenium (Se), Zn, Antimony (Sb) (Başgel & Erdemoğlu), Arsenic (As),
Cr, Cobalt (Co), Lead (Pb), Lithium (Li), Molybdenum (Mo), Nickel (Ni), Titanium
(Ti) (Turco et al.), Tin (Sn) (Al-Snafi), cadmium (Cd) and Aluminium (Al) (Ficsor
et al., 2013; Soni et al., 2014).
472 S. A. Mahesar et al.

4.1 The Plant

The fig plant can reach a height of 18–20 feet. The stem is medium in size, with a
smattering of branches. Roots are not adventitious and are found underground. The
milky white latex found in the stems and leaves contains ficin, a protein-degrading
enzyme. The fig tree leaves are primarily green and petiolated, with curling borders
and a hairy surface (Badgujar & Mahajan, 2009). The green fruit turns yellowish or
red-black as it reaches maturity. The fruit and leaves of F. carica have long been
used to treat throat ailments and as a sedative, laxative, emollientantitussive, resol-
vent, emmenagogue, constipation, hemorrhoid, and high cholesterol (Ghazi et al.,
2012). The traditional use of fig leaves in treating diseases such as vitiligo, diabetes,
coughs, asthma, constipation, and gingivitis has also been reported (Mohan et al.,
2007). Furthermore, roots used to cure leukoderma and ringworms and fruits have
antipyretic, purgative, and aphrodisiac effects and have been demonstrated to be
beneficial in paralysis and inflammations (Rahmani & Aldebasi, 2017). Fig leaves
contain the minerals are Mg, Mn, Fe, Ca, Cu, Na, K, and P, as shown in (Table 20.2).

4.2 Seeds

The fig is a fruit eaten with or without its peel and contains many seeds in its jelly-
like flesh. The seeds are one of the most appealing portions of whole figs, and they
impact both their nutritional value and their health consequences. The seeds are

Table 20.2 Mineral composition of fig leaves

Fig leaves Elements (mg/100g) Reported studies
(Ghazi et al., 2012) (Essid et al., 2015)
Mg 307.9–396.4 160.9–416.3
Mn 21.9–23.1 0.84–6.09
Fe 66.3–75.5 17.2–48.2
Ca 1398.1–2551.3 169.2–779.1
Na – 45.76–391.6
K 117.67–496.71 379.05–1412.8
P – 102.1–323.8
20 Fig Minerals 473

Table 20.3 Mineral composition of fig seeds

Fig seeds Elements (mg/100g) Reported studies
(Nakilcioğlu-Taş, 2019) (Duman et al., 2018)
Mg 4.64–5.37 –
Mn 0.77–0.803 –
Fe 1.83–2.26 0.19
Ca 18.57–19.04 –
Cu 0.525–0.534 0.619
Na 6.945–7.082 –
K 16.67–17.28 –
P 17.83–18.62 –
Zn – 0.0219
As – <0
Cr – 0.0785
Co – 0.0056
Ni – 0.158
Al – 0.0001

smaller, edible, and come in various sizes such as giant, medium, small, or minute,
with 30–1600 seeds per fruit (Badgujar et al., 2014; Lansky et al., 2008). For this
reason, the nutritional composition of figs is thought to be influenced mainly by the
fig seeds (Nakilcioğlu-Taş, 2019). Table 20.3 shows the mineral composition of
fig seeds.

4.3 The Ripe Fruits

The ripe fruits can be consumed fresh, dried, or preserved in other ways (Kamiloglu
& Capanoglu, 2015). If refrigerated, fully ripened fresh figs have 2–3 days of shelf
life. In the sun, they require 4–5 days to dry, and in a dehydrator, they take 10–12 h
to dry. Dried figs are a good source of nutrients, particularly essential minerals and
energy, and can be stored for six to eight months. Table 20.4 shows the mineral
composition of fig fruit.
474 S. A. Mahesar et al.

Table 20.4 Mineral composition of fig fruit

Elements Reported studies
(mg/100g) (Aljane & (M. N. (Soni (Mahmoudi (Turco
Ferchichi, Khan et al., et al., et al., 2018) et al.,
2009) 2011) 2014) 2020)
Fig Mg 47.1–99.4 110.5–202.4 67.9 – 0.084–
Fruit 0.128
Mn 0.21–0.43 – 0.475 – 0.353–1.4
Fe 0.08–1.17 5.69–10.1 2.95 – 1.27–2.93
Ca 167.6–403.2 78.7–132.8 154.6 – 0.146–
0.449
Cu 0.17–0.43 0.25–0.42 0.502 13.9–125.4 0.425–
0.812
Na 38.20–117.0 5.58–17.84 – 4.63–14.12 0.045–0.07
K 688.8–1047 382.4–611.6 – 165.3–266.7 0.35–0.812
P – 31.91–76.96 36.575 7.62–16.5 –
Zn 0.39–0.69 0.32–0.62 0.987 – 0.523–1.15
Sb – – 0.0298 – –
As – – 0.1669 – 0.006–
0.014
Cr – – 0.147 – 0.003–
0.008
Co – – 0.0032 – –
Pb – – 0.0680 – 0–0.006
Li – – 0.0025 – 0.003–
0.008
Mo – – 0.0026 – –
Ni – – 0.118 – 0.014–
0.054
Se – – 0.079 – –
Ti – – 0.037 – –
Sn – – 0.133 – –
Cd – – 0.0003 – 0–0.006

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Chapter 21
Bioactive Compounds of Fig (Ficus carica)

Senem Kamiloglu and Banu Akgun

1 Introduction

The common fig (Ficus carica L.), belonging to the Moraceae family, is one of the
most abundant fruits in the Mediterranean diet, promoting health by preventing-
pathophysiological conditions related to chronic diseases (Kamiloglu & Capanoglu,
2013). Traditionally, Ficus carica has been used to treat disorders linked with the
endocrine system (e.g., diabetes), respiratory system (e.g., asthma, cough, and liver
diseases), gastrointestinal tract (e.g., vomiting and ulcer), and reproductive system
(e.g., menstruation pain), as well as infectious diseases (e.g., scabies, gonorrhea,
and skin disease) (Badgujar et al., 2014). These health-promoting effects of figs are
associated with the presence of bioactive compounds.
The fig cultivation originated in the East Mediterranean region, which was later
expanded into the West Mediterranean area (Teruel-Andreu et al., 2021). Today,
with an annual production of 310 thousand tons, Türkiye is the world’s leading fig
producing country, accounting for approximately 25% of the world’s total produc-
tion. Other major fig producing countries include Egypt, Morocco, Iran, Algeria,
and Spain. Therefore, the Mediterranean basin and the Near East remain crucial for

S. Kamiloglu (*)
Department of Food Engineering, Faculty of Agriculture, Bursa Uludag University,
Gorukle, Bursa, Türkiye
Science and Technology Application and Research Center (BITUAM), Bursa Uludag
University, Gorukle, Bursa, Türkiye
e-mail: [email protected]
B. Akgun
Food Authenticity and Allergen Department, Central Research Institute of Food and Feed
Control, Bursa, Türkiye
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 479
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_21
480 S. Kamiloglu and B. Akgun

fig production. Although not listed among the top fig-producing countries, Australia
and Netherlands are reported to be the largest exporters of this crop, with a com-
bined value of over US$30 million in 2019 (FAOSTAT, 2021).
The peel color of fig fruits ranges from green to dark purple. Although figs are
often consumed after removing the peels, they can also be consumed together with
the peel (Solomon et al., 2006). Fresh figs have a short post-harvest life ranging
between a week to 10 days. However, the combination of a cooled and carbon
dioxide-enriched environment enhances the shelf life of the fruit by up to a month.
Figs are also often consumed as dried fruits that can be stored for 6–8 months,
ensuring proper preservation (Slatnar et al., 2011; Veberic et al., 2008). Some other
processed fig products include jam, marmalade, or paste utilized in various bakery
products (Saif et al., 2020).
Phytochemical studies on figs and their processed products revealed the presence
of several bioactive compounds, including polyphenols, carotenoids, vitamins,
organic acids, triterpenoids, phytosterols, and fatty acids. The concentration of these
bioactive compounds is highly dependent on the fig variety, cultivation technique,
environmental conditions, and processing parameters, among others. In this chapter,
we provided an overview of the findings in the literature on the bioactive com-
pounds of figs. The levels of polyphenols, carotenoids, vitamins, organic acids, tri-
terpenoids, phytosterols, and fatty acids in figs were discussed in detail, along with
the parameters that influence the levels of these compounds.

2 Polyphenols

Polyphenols are secondary plant metabolites that possess antioxidant, antimicrobial,

antiviral, and anti-inflammatory properties. According to epidemiological studies
and related meta-analyses, long-term intake of diets rich in plant polyphenols pro-
tects against the development of chronic diseases such as cardiovascular diseases,
diabetes, certain types of cancers, and neurodegenerative diseases. Polyphenols are
classified into different groups based on the number of phenol rings they include and
the structural components (Ignat et al., 2011; Pandey & Rizvi, 2009). According to
the findings in the literature, fig fruits are rich sources of polyphenols. However, the
two most well-known sources of polyphenols, red wine and tea, contain lower levels
of polyphenols than figs (Mawa et al., 2013; Vallejo et al., 2012). The major poly-
phenol groups present in fig fruits include flavonoids, phenolic acids, and coumarins
(Fig. 21.1), which are discussed in detail in the following sections.

2.1 Flavonoids

Flavonoids are low molecular weight molecules with 15 carbon atoms organized in
a C6-C3-C6 pattern. In the flavonoid structure, two aromatic rings are connected by
a 3-carbon bridge, which is typically in the form of a heterocyclic ring. Different
21 Bioactive Compounds of Fig (Ficus carica) 481

Polyphenols in figs (Ficus carica L.)

Flavonoids Phenolic Acids Coumarins

Psoralen

Chlorogenic acid

Anthocyanins Flavonols Flavanols Flavones

Cyanidin 3-rutinoside Rutin Catechin Apigenin

Fig. 21.1 Major classes of polyphenols in figs (Ficus carica L.)

subclasses arise from variations in the heterocyclic ring’s substitution pattern

(Balasundram et al., 2006). Among these subclasses, anthocyanins, flavonols, flava-
nols, and flavones are the ones that are reported to be present in figs.

2.1.1 Anthocyanins

Anthocyanins are water-soluble pigments responsible for the blue, purple, and red
color of fruits (particularly berries), vegetables, legumes, roots, and cereals. They
belong to the group of compounds known as flavonoids and are found as glycosides
of their respective aglycones, called anthocyanidins. Cyanidin, delphinidin, malvi-
din, pelargonidin, peonidin, and petunidin are major anthocyanidins present in
foods (Kamiloglu et al., 2015). As anthocyanins possess antioxidant, anti-
inflammatory, antiatherogenic, and anti-apoptotic properties, their consumption is
suggested to treat non-communicable diseases, including cardiovascular diseases,
obesity, diabetes, and neurodegenerative diseases, to modulation bacteria and bone
metabolism (Speer et al., 2020).
The anthocyanins present in fig fruits were quantified using spectrophotometric
or chromatographic methods. The pH differential method is the most employed
technique to determine the total anthocyanin content of figs. Cyanidin 3-rutinoside
and cyanidin 3-glucoside are commonly used as a reference standard to express the
results of total anthocyanin content of figs (Aljane et al., 2020; Çalişkan & Polat,
2011; Ercisli et al., 2012; Harzallah et al., 2016; Hssaini et al., 2019; Hssaini et al.,
2020a; Martínez-García et al., 2013; Meziant et al., 2018, 2021; Petkova et al.,
2019; Rodov et al., 2012; Saki et al., 2019; Solomon et al., 2006; Viuda-Martos
et al., 2015) (Table 21.1). Although it is pretty difficult to compare the results
Table 21.1 Anthocyanin content of figs (Ficus carica L.)
Variety/Origin Skin color Form/Part Compound(s) Content References
482

Mission/Israel Black Fresh Total anthocyanins 10.9 mg C3G/100 g Solomon et al.

Cyanidin 3-rutinoside n/a (2006)
Cyanidin 3-glucoside n/a
Chechick, Brown Türkiye, Purple, red Fresh Total anthocyanins 0.3–1.8 mg C3G/100 g
and Bursa/Israel
Mattalona/Italy Black Peel Cyanidin 3-rutinoside 901.03 mg/kg FW del Caro and
Cyanidin 3-glucoside 28.24 mg/kg FW Piga (2008)
Pulp Cyanidin 3-rutinoside 4.92 mg/kg FW
San Pietro/Italy Green Pulp Cyanidin 3-rutinoside 17.60 mg/kg FW
Colar, Granilla and Cuello Dark purple Skin Cyanidin 3-rutinoside 15.42–78.32 μg/g FW Dueñas et al.
de Dama/Salamanca, Spain; Cyanidin 3-glucoside 1.51–15.38 μg/g FW (2008)
Bursa Siyahi/Türkiye Cyanidin 3,5-diglucoside 1.99–5.23 μg/g FW
Pelargonidin 3-rutinoside 0.47–3.52 μg/g FW
Cyanidin 3-malonylglucoside 0.67–3.51 μg/g FW
Cyanidin 3-malonylglycosyl-5-glucoside 0.53–1.39 μg/g FW
Carboxypyrano-cyanidin 3-rutinoside 0.48–1.26 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-rutinoside 0.53–1.23 μg/g FW
Cyanidin 3-rutinoside dimer 0.44–0.93 μg/g FW
Pelargonidin 3-glucoside 0.29–0.68 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-glucoside 0.38–0.48 μg/g FW
(Epi)catechin-(4→8)-pelargonidin 0.20–0.24 μg/g FW
3-rutinoside
Pulp Cyanidin 3-rutinoside 3.58–10.16 μg/g FW
Cyanidin 3-glucoside 0.51–2.15 μg/g FW
Carboxypyrano-cyanidin 3-rutinoside 0.05–0.94 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-rutinoside 0.06–0.59 μg/g FW
Cyanidin 3,5-diglucoside 0.11–0.57 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-glucoside 0.05–0.13 μg/g FW
Cyanidin 3-malonylglycosyl-5-glucoside 0.04–0.09 μg/g FW
Cyanidin 3-malonylglucoside 0.05–0.09 μg/g FW
Pelargonidin 3-glucoside 0.01–0.09 μg/g FW
S. Kamiloglu and B. Akgun

Pelargonidin 3-rutinoside 0.04–0.08 μg/g FW

Peonidin 3-rutinoside 0.02–0.06 μg/g FW
Cyanidin 3-rutinoside dimer 0.08 μg/g FW
Variety/Origin Skin color Form/Part Compound(s) Content References
Cuello de Dama/Salamanca, Green Pulp Cyanidin 3-rutinoside 1.04 μg/g FW Dueñas et al.
Spain Cyanidin 3-glucoside 0.19 μg/g FW (2008)
Cyanidin 3,5-diglucoside 0.09 μg/g FW
Carboxypyrano-cyanidin 3-rutinoside 0.04 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-glucoside 0.03 μg/g FW
(Epi)catechin-(4→8)-cyanidin 3-rutinoside 0.02 μg/g FW
Cyanidin 3-malonylglycosyl-5-glucoside 0.02 μg/g FW
Cyanidin 3-malonylglucoside 0.02 μg/g FW
Pelargonidin 3-rutinoside 0.02 μg/g FW
76 local Mediterranean fig Green Fresh Total anthocyanins 0.0–41.3 μg C3R/g FW Çalişkan and
accessions/Hatay, Türkiye Yellow Fresh Total anthocyanins 0.6–29.7 μg C3R/g FW Polat (2011)
Brown Fresh Total anthocyanins 2.0–49.6 μg C3R/g FW
Purple Fresh Total anthocyanins 2.5–46.5 μg C3R/g FW
Black Fresh Total anthocyanins 32.3–356.0 μg C3R/g FW
Bela petrovka/Slovenia n/a Fresh Cyanidin 3-rutinoside 0.21–0.62 mg/100 g Slatnar et al.
21 Bioactive Compounds of Fig (Ficus carica)

Oven- Cyanidin 3-rutinoside 0.12–0.31 mg/100 g (2011)

dried
Sun-dried Cyanidin 3-rutinoside 0.12–0.26 mg/100 g
24 local cultivars and Bursa Black Fresh Total anthocyanins 31–42 mg C3R/100 g Ercisli et al.
Siyahi/Coruh valley, Light purple Fresh Total anthocyanins 8–10 mg C3R/100 g (2012)
Türkiye Purple Fresh Total anthocyanins 13–20 mg C3R/100 g
Dark purple Fresh Total anthocyanins 32 mg C3R/100 g
Brown Türkiye/Bet Dagan, Light Pulse Total anthocyanins 0.1–4 mg/100 g Rodov et al.
Israel green-purple lighted (2012)
18 cultivars/Alicante, n/a Skin Cyanidin 3-rutinoside 16.9–108.9 mg/100 g FW Vallejo et al.
Spain Cyanidin 3-glucoside 0.6–11.1 mg/100 g FW (2012)
Pulp Cyanidin 3-rutinoside 1.0–9.5 mg/100 g FW
Bursa siyahi, Morguz and Green-purple Fresh Cyanidin 3-rutinoside 21.27–91.23 mg/100 g FW Yemiş et al.
Yesilguz/Türkiye Cyanidin 3-glucoside 2.61–7.07 mg/100 g FW (2012)
Cyanidin 3,5-diglucoside 2.91–6.24 mg/100 g FW
483

(continued)
Table 21.1 (continued)
484

Variety/Origin Skin color Form/Part Compound(s) Content References

Mission/Mexico n/a Fresh Total anthocyanins 1.12–1.42 mg/100 g Martínez-García
et al. (2013)
Bouholi and Zidi/Tunisia Dark purple Fresh Cyanidin 3-rutinoside 7.6–13.9 mg/100 g FW Trad et al.
Cyanidin 3-glucoside 0.2–0.4 mg/100 g FW (2014)
Thgagli, Bidhi, and Yellow-green Fresh Cyanidin 3-rutinoside 3.5–4.2 mg/100 g FW
Khendri/Tunisia Cyanidin 3-glucoside 0.1–0.7 mg/100 g FW
Tounsi and Temri/Sfax, Purple Dried Cyanidin 3-rutinoside n/a Ammar et al.
Tunisia Cyanidin 3,5-diglucoside n/a (2015)
Sarilop/Aydin, Türkiye Yellow Skin, Pulp Cyanidin 3-rutinoside 0.1–2.2 mg/100 g DW Kamiloglu and
Cyanidin 3-glucoside 0.1 mg/100 g DW Capanoglu
Bursa siyahi/Bursa, Türkiye Purple Skin, Pulp Cyanidin 3-rutinoside 18–180 mg/100 g DW (2015)
Cyanidin 3-glucoside 2.8–23 mg/100 g DW
Dried Cyanidin 3-rutinoside 1.5 mg/100 g DW
Cyanidin 3-glucoside 0.1 mg/100 g DW
Colar and Cuello de dama/ Purple Peel Total anthocyanins 6.21–6.63 mg C3G/g Viuda-Martos
Spain et al. (2015)
Tounsi/Sfax, Tunisia n/a Dried Cyanidin 3-rutinoside n/a Belguith-
Hadriche et al.
(2016)
Bidhi, Hamri and Kohli/ Green, Purple, Peel, Pulp Total anthocyanins 144.62–3330.84 mg C3G/100 g Harzallah et al.
Bekalta, Tunisia Black FW (2016)
10 different cultivars/ Black, Green, Fresh Cyanidin 3-rutinoside 0.4–116.2 mg/100 g DW Wojdyło et al.
Alicante, Spain Greenish Pelargonidin 3-rutinoside 0.0–3.8 mg/100 g DW (2016)
yellow, Cyanidin 3,5-diglucoside 0.0–1.9 mg/100 g DW
Brown-green,
Intense purple
S. Kamiloglu and B. Akgun
Variety/Origin Skin color Form/Part Compound(s) Content References
De Rey, Cuello Dama Brown, Dark Skin Cyanidin 3-rutinoside 12.19–97.4 mg/100 g Pereira et al.
Negro, Colar Elche, San purple Pelargonidin 3-rutinoside 0.09–5.64 mg/100 g (2017)
Antonio, and Brown Cyanidin 3,5-diglucoside 0.12–6.39 mg/100 g
Türkiye cultivars/Guadajira, Pulp Cyanidin 3-rutinoside 1.21–5.92 mg/100 g
Badajoz, Spain Cyanidin 3,5-diglucoside 0.01–0.66 mg/100 g
Pelargonidin 3-rutinoside 0.01–0.26 mg/100 g
Azenjer/Bejaia, Algeria Black Peel Total anthocyanins 175.76–326.82 mg/100 g DW Meziant et al.
(2018)
Sabz/Estahban, Iran Yellow Fresh Cyanidin 3-malonyl-glucoside 0.0–0.65 mg/g Sedaghat and
Cyanidin 3-rutinoside 0.25–0.43 mg/g Rahemi (2018)
Pelargonidin 3-glucoside 0.0–0.40 mg/g
Cyanidin 3-glucoside 0.0–0.28 mg/g
135 genotypes/Ain- Green-yellow, Fresh Total anthocyanins 0.41–57.45 mg/100 g DW Hssaini et al.
Taoujdate, Morocco purple (2019)
Croatian Yellow Giant/ Yellow-green Fresh Total anthocyanins 0.8 mg C3G/100 g Petkova et al.
21 Bioactive Compounds of Fig (Ficus carica)

Plovdiv, Bulgaria Jam Total anthocyanins 0.4 mg C3G/100 g (2019)

Unknown/Durango, Mexico n/a Uncoated Cyanidin 3-rutinoside 53.1–116.93 mg/100 g DW Reyes-Avalos
Cyanidin 3-glucoside 4.80–13.00 mg/100 g DW et al. (2019)
Coated Cyanidin 3-rutinoside 19.43–67.38 mg/100 g DW
Cyanidin 3-glucoside 2.79–6.73 mg/100 g DW
Siah/Lorestan, Iran Deep purple Uncoated Total anthocyanins ~ 1.7–3.2 mg/100 g Saki et al.
Coated Total anthocyanins ~ 3.0–3.8 mg/100 g (2019)
27 ecotypes/Medenine, Green Fresh Total anthocyanins 1.61–11.67 mg/100 g FW Aljane et al.
Tunisia yellowish, (2020)
Green, Red
greenish, Brown
purplish, Purple
greenish, Purple
blackish
(continued)
485
Table 21.1 (continued)
486

Variety/Origin Skin color Form/Part Compound(s) Content References

Five local cultivars/ Reddish purple Fresh Total anthocyanins 3.1–24.8 mg C3R/100 g DW Hssaini et al.
Morocco, six introduced (2020a, b)
cultivars from USA, Italy
and Spain
Brown Türkiye/Israel Green-purple Untreated Cyanidin 3-glucoside 1.38–5.47 μg/g FW Lama et al.
Cyanidin 3-rutinoside 0–2.45 μg/g FW (2020)
ABA- Cyanidin 3-glucoside 14.13–1.66 μg/g FW
treated Cyanidin 3-rutinoside 2.55–10.39 μg/g FW
ETP- Cyanidin 3-glucoside 10.79–12.35 μg/g FW
treated Cyanidin 3-rutinoside 2.85–4.84 μg/g FW
Bpjihon, Masui Dauphine, Brown, purple, Peel Cyanidin 3-glucoside 1.14–58.54 mg/100 g FW Zhang et al.
Wild-type, Japanese purple red, Cyanidin 3,5-diglucoside 0–30.73 mg/100 g FW (2020)
fruit purple-brown Cyanidin 3-rutinoside 0–5.60 mg/100 g FW
Pulp Cyanidin 3-glucoside 7.72–42.11 mg/100 g FW
Cyanidin 3,5-diglucoside 0–14.90 mg/100 g FW
Cyanidin 3-rutinoside 0.40–3.01 mg/100 g FW
Vasilika, Mission and MM/ Green, purple Peel Cyanidin 3-rutinoside 0.01–910.6 mg/100 g FW Karantzi et al.
Markopoulo, Greece Cyanidin 3-glucoside 0.01–38.2 mg/100 g FW (2021)
Flesh Cyanidin 3-rutinoside 14.2–212.2 mg/100 g FW
Cyanidin 3-glucoside 0.6–16.7 mg/100 g FW
Bakor Noir, Bouankik, Greenish- Fresh Total anthocyanins 7.21–62.22 mg C3R/mL Meziant et al.
Azenjer, Tazegaght/Algeria purple, (2021)
Reddish-purple,
black,
Reddish-violet
ABA abscisic acid, C3G cyanidin 3-glucoside, C3R cyanidin 3-rutinoside, DW dry weight, ETP ethephon, FW fresh weight, n/a not available
S. Kamiloglu and B. Akgun
21 Bioactive Compounds of Fig (Ficus carica) 487

expressed using different standards, the outcomes obtained using the spectrophoto-
metric pH differential method revealed that the total anthocyanin content in brown,
purple and black fresh figs was higher than that of green and yellow figs. This obser-
vation is related to the anthocyanins in dark-colored figs primarily accumulated in
peels. Food processing also affected the total anthocyanin content. According to the
study carried out by Petkova et al. (2019), the total anthocyanin content of yellow-
green figs from Bulgaria was reduced by 50% after jam processing. Moreover,
freezing diminished the anthocyanins in figs. In another study (Saki et al., 2019),
chitosan and thymol-coated purple figs from Iran exhibited higher total anthocyanin
content than uncoated figs. Therefore, the authors proposed coating as an efficient
method for quality maintenance and shelf life extension of fresh fig fruits.
High-performance liquid chromatography (HPLC) and ultra-performance liquid
chromatography (UPLC) are the most common instruments used to separate, iden-
tify, and quantify anthocyanins. The major anthocyanins detected in figs were
cyanidin- based with different sugar moieties (Ammar et al., 2015; Belguith-
Hadriche et al., 2016; del Caro & Piga, 2008; Dueñas et al., 2008; Kamiloglu &
Capanoglu, 2015; Karantzi et al., 2021; Lama et al., 2020; Pereira et al., 2017;
Reyes-Avalos et al., 2019; Sedaghat & Rahemi, 2018; Slatnar et al., 2011; Solomon
et al., 2006; Trad et al., 2014; Vallejo et al., 2012; Wojdyło et al., 2016; Yemiş et al.,
2012; Zhang et al., 2020) (Table 21.1). In particular, cyanidin 3-rutinoside and cyan-
idin 3-glucoside were detected as the main anthocyanins in figs with different skin
colors. Besides cyanidin-based anthocyanins, pelargonidin 3-rutinoside, pelargoni-
din 3-glucoside, and peonidin 3-rutinoside were also identified in different figs cul-
tivars from Spain and Iran (Dueñas et al., 2008; Pereira et al., 2017; Sedaghat &
Rahemi, 2018; Wojdyło et al., 2016). Similar to the results obtained using the spec-
trophotometric pH differential method, higher concentrations of anthocyanins were
detected in dark-colored fruits than in light-colored figs. Moreover, peels of purple
fıgs are found to contain higher levels of anthocyanins than pulps (del Caro & Piga,
2008; Dueñas et al., 2008; Kamiloglu & Capanoglu, 2015; Karantzi et al., 2021;
Pereira et al., 2017; Vallejo et al., 2012; Zhang et al., 2020).
Anthocyanins are prone to chemical degradation by processing and storage con-
ditions, such as temperature, light, pH, and oxygen (Oancea, 2021). In general,
thermal processing results in reduced anthocyanin content and bioaccessibility. On
the other hand, it has been indicated that thermal treatments damage cell walls,
liberating the bound anthocyanins and thus enabling more accessible compounds
for absorption (Eker et al., 2020). Accodingly, Slatnar et al. (2011) reported that
both sun- and oven-drying caused degradation in cyanidin 3-rutinoside content of
figs from Slovenia (26–61%). Similarly, significant decreases (96–98%) were
observed in cyanidin 3-rutinoside and cyanidin 3-glucoside contents of Turkish figs
after sun-drying (Kamiloglu & Capanoglu, 2015). Furthermore, no anthocyanins
were detected in the dialyzed fraction after in vitro gastrointestinal digestion of sun-
dried figs (Kamiloglu & Capanoglu, 2013). In another study (Reyes-Avalos et al.,
2019), the effect of alginate-chitosan coating on the anthocyanin content of fresh
figs was investigated. The results revealed lower contents of cyanidin 3-rutinoside
and cyanidin 3-glucoside in coated figs than in uncoated ones. Reduced anthocyanin
488 S. Kamiloglu and B. Akgun

levels in the coated figs could be related to the internal atmosphere modification, as
it has been reported that the biochemical reactions involved in the synthesis of
anthocyanins can be regulated by a modified atmosphere (Reyes-Avalos et al.,
2019). In addition, a recent study (Lama et al., 2020) showed that abscisic acid pre-
treatment could initiate the biosynthesis of cyanidin 3-rutinoside and cyanidin
3-glucoside enhancing the nutritional value and the color of fig fruits. Overall, the
studies conducted in the literature pointed out that anthocyanin composition and
concentration in fig fruits depends on the variety, processing conditions, as well as
pre- and post-harvest treatments.

2.1.2 Flavonols

Flavonols are the most common subgroup of the flavonoids. Quercetin, kaempferol,
myricetin, and isorhamnetin are the major dietary flavonol aglycones, often occur-
ring as O-glycosides in dietary sources. Flavonols possess several important bio-
logical activities, including antioxidant, anti-inflammatory, antiangiogenic,
hypolipidemic, neuroprotective, and anticancer effects (Kamiloglu et al., 2020).
Many studies carried out in the literature identified rutin (quercetin 3-rutinoside)
as the major flavonol present in figs fruits (Feng et al., 2015; Karantzi et al., 2021;
Veberic et al., 2008; Viuda-Martos et al., 2015) and leaves (Ladhari et al., 2020; Li
et al., 2021; Petruccelli et al., 2018; Qin et al., 2015; Takahashi et al., 2017; Teixeira
et al., 2006). According to Wojdyło et al. (2016), the content of rutin corresponded
to 55–80% of total flavonols identified in fig fruits. Besides rutin, quercetin, iso-
quercetin (quercetin 3-glucoside) (Gaaliche et al., 2019), quercetin 3-(6″-malonyl)
glucoside, quercetin deoxyhexoside dihexoside, quercetin dihexoside, quercetin
dideoxyhexoside hexoside (Ammar et al., 2015), quercetagetin 7-glucoside, kaemp-
ferol 7-glucoside (Francini et al., 2021), kaempferol 3-glucoside (Slatnar et al.,
2011), kaempferol 3-rutinoside (Kamiloglu & Capanoglu, 2015; Wojdyło et al.,
2016), kaempferol deoxyhexosyl-hexoside and taxifolin hexoside (Palmeira et al.,
2019) were also detected in figs.
In general, higher concentrations of flavonols were found in the fig peels than in
pulps (Palmeira et al., 2019). Accordingly, black figs contained higher levels of
flavonols than green or yellow figs (del Caro & Piga, 2008; Kamiloglu & Capanoglu,
2015). Sedaghat and Rahemi (2018) reported that the concentration of flavonols,
including rutin, isoquercetin, and kaempferol 3-glucoside, increases as figs mature.
Potassium application also increased quercetin levels significantly (from 0.19 to
4.29 mg/g DW). However, potassium treatment also reduced the rutin content sig-
nificantly (two times compared to the control) (Gaaliche et al., 2019). Drying is
another parameter that influences the flavonol concentration in figs. According to
Slatnar et al. (2011), statistically higher rutin contents, isoquercetin, and kaempferol
3-glucoside were determined in figs dried in the drying oven. On the other hand,
Kamiloglu and Capanoglu (2015) showed that the sun-drying process caused
decreases in all flavonols of yellow figs (24–50%). In contrast, rutin and kaempferol
rutinoside contents increased in sun-dried purple figs (16–200%). Li et al. (2021)
21 Bioactive Compounds of Fig (Ficus carica) 489

indicated that flavonols from fig leaves possessed significant antioxidant activity.
Furthermore, in a study investigating the bioactivity of a fig variety from the
Shandong province of China, quercetin was identified as the major antiproliferative
compound along with trace amounts of rutin (Mustafa et al., 2021).

2.1.3 Flavanols

Epicatechin and catechin are the major flavanols present abundantly in fruits, cocoa,
red wine, and green tea (Hackman et al., 2008). The main physiological functions
and regulating mechanisms of flavanols consist of antioxidant, anti-inflammatory,
antidiabetic, anticancer, cardioprotective, and neuroprotective activities and
enhanced skeletal muscle performance (Li et al., 2019).
From the group of flavanols, both catechin and epicatechin have been detected in
fig fruits (Francini et al., 2021). Vallejo et al. (2012) also suggested the presence of
additional polymeric procyanidins, up to tetramers, in fig fruits. The majority of the
studies conducted in the literature identified catechin as the main flavanol (up to
37.8 mg/100 g FW) (Feng et al., 2015; Kamiloglu & Capanoglu, 2015; Pande &
Akoh, 2010; Tsanova-Savova et al., 2005; Veberic et al., 2008; Viuda-Martos et al.,
2015). On the other hand, Slatnar et al. (2011) reported epicatechin as the predomi-
nant compound (7.11–8.67 mg/100 g). In general, peels contained higher flavanols
compared to pulps (Pereira et al., 2017; Viuda-Martos et al., 2015). However,
Kamiloglu and Capanoglu (2015) indicated that although catechin was primarily
located in the fruit skin for yellow figs, purple figs pulp contained higher compound
levels. These differences may be cultivar-specific or related to the agroecological
conditions of these studies (Wojdyło et al., 2016). Pereira et al. (2017) and Sedaghat
and Rahemi (2018) found that the levels of flavanols increased during ripening.
Effects of drying on the flavanol content of fig fruits varied. While Slatnar et al.
(2011) observed an increase in epicatechin contents in Slovenian figs as a result of
sun-drying (from 7.8 mg/100 g FW in fresh figs to 25.4 mg/100 g DW in dried figs),
Kamiloglu and Capanoglu (2015) reported decreased levels for both catechin
(35–45%) and epicatechin (45–68%) after sun-drying of Turkish figs.

2.1.4 Flavones

Apigenin and luteolin are the main flavone aglycones present in fruits, vegetables,
herbs, and beverages. Flavones have demonstrated various potential health-
beneficial activities in human trials and animal studies (Hostetler et al., 2017). For
example, human trials reported improved effects on Alzheimer’s disease, osteoar-
thritis, insomnia, depression, and anxiety. Similarly, animal models suggested ther-
apeutic effects on diabetes, cancer, Alzheimer’s disease, depression, and amnesia
(Salehi et al., 2019).
Flavones primarily present in fig fruits include apigenin, luteolin, and their C-
and O-glycosidic forms. In particular, Vallejo et al. (2012) reported the presence of
490 S. Kamiloglu and B. Akgun

luteolin 6C-hexose-8C-pentose and apigenin-rutinoside in figs from Spain, with the

latter being the most abundant flavone in fruit peel (0.8–2.5 mg/100 g FW).
Similarly, Ammar et al. (2015) and Soltana et al. (2018) detected apigenin, luteolin,
and their C-glycosidic forms and luteolin 7-O-glucoside in different fig varieties
from Tunisia. On the other hand, in another study investigating the flavones in pur-
ple figs from Spain, while apigenin and luteolin were detected both in peel and pulp,
luteolin 7-O-glucoside was only present in the peel (Viuda-Martos et al., 2015).
Palmeira et al. (2019) also determined C-glycosylated apigenin derivatives only in
the peel of Portuguese figs. Apigenin C-hexoside-pentoside ranging from 0.53 to
24.4 mg/100 g DW was the only flavone detected according to the study of Wojdyło
et al. (2016). Besides the fig fruits, fig leaves also contained apigenin, luteolin, and
their glycosides (Ammar et al., 2015; Mustafa et al., 2021; Zhao et al., 2021). The
drying process changed the flavone content of figs. For purple Turkish figs, the api-
genin content was increased significantly (68%) due to drying, whereas for yellow
figs, there was no statistically significant difference (Kamiloglu & Capanoglu, 2015).

2.2 Phenolic Acids

Phenolic acids can be categorized as hydroxycinnamic and hydroxybenzoic acids,

having different molecular skeletons based on cinnamic and benzoic acids, respec-
tively (Song et al., 2020). Hydroxycinnamic acids include caffeic, ferulic, p-
coumaric, and sinapic acids, while gallic, protocatechuic, vanillic, and syringic
acids are the most common hydroxybenzoic acids. In nature, hydroxycinnamic
acids are more prevalent than hydroxybenzoic acids and commonly occur in conju-
gated forms. Various fruits, vegetables, spices, grains, and beverages contain pheno-
lic acids (Rashmi & Negi, 2020). Chlorogenic acid is also one of the most common
hydroxycinnamic acids with quinic acid that exerts various therapeutic functions
such as antioxidant, antimicrobial, anti-inflammatory, cardioprotective, hepatopro-
tective, and central nervous system stimulator properties (Naveed et al., 2018).
The majority of the studies investigating the phenolic acid content of figs reported
chlorogenic acid as the main phenolic acid in the fruits (del Caro & Piga, 2008;
Feng et al., 2015; Slatnar et al., 2011; Vallejo et al., 2012; Wojdyło et al., 2016) and
leaves (Qin et al., 2015; Teixeira et al., 2006). Besides chlorogenic acid, gallic acid
(Qin et al., 2015; Reyes-Avalos et al., 2019), syringic acid (Veberic et al., 2008),
ellagic acid (Pereira et al., 2017), p-coumaric acid (Kamiloglu & Capanoglu, 2015)
and protocatechuic acid (Viuda-Martos et al., 2015) were also detected in figs.
Ammar et al. (2015) performed a comprehensive identification of phenolics in
Tunisian figs and reported the presence of 24 hydroxycinnamic acids, which were
derivatives of coumaric, caffeic, ferulic, and sinapic acids. Chlorogenic acid and its
other three isomers were also found.
Several factors are reported to influence the phenolic acid content in figs. For
instance, skin color is one of the parameters affecting the levels of phenolic acids.
Accordingly, purple figs were richer in phenolic acids than fellow figs (Kamiloglu
21 Bioactive Compounds of Fig (Ficus carica) 491

& Capanoglu, 2015). Similarly, the fruit part is another factor that affects the phe-
nolic acid content. In general, higher phenolic acids were found in the fig peels than
in pulps (Pereira et al., 2017; Vallejo et al., 2012). On the other hand, Kamiloglu and
Capanoglu (2015) reported higher content of gallic acid in pulp compared to skin.
Likewise, Viuda-Martos et al. (2015) also determined higher amounts of protocat-
echuic acid in pulp than in peel. According to Sedaghat and Rahemi (2018), the
chlorogenic acid content of figs increased during ripening. Furthermore,
Pourghayoumi et al. (2012) reported that levels of chlorogenic acid in figs were
affected by the pollen sources, female cultivars, and their interaction. Another study
showed that potassium treatment significantly enhanced the chlorogenic acid con-
tent (Gaaliche et al., 2019).
The drying process had different impacts on the phenolic acid content of fig
fruits. According to Slatnar et al. (2011), statistically higher amounts of chlorogenic
acid were measured in figs dried in the drying oven (1.3–4.9 mg/100 g and
3.4–32.4 mg/100 g in fresh and dried figs, respectively). On the other hand,
Kamiloglu and Capanoglu (2015) indicated that drying reduced the chlorogenic
acid and p-coumaric acid contents (34–87%), while an increase of 67% was
observed in gallic acid content. A study investigating the effect of alginate-chitosan
coating on the phenolic acid content of figs (Reyes-Avalos et al., 2019) revealed that
coated figs exhibited higher gallic acid content (1.7 mg/100 g DW) compared to
uncoated figs. (2.8 mg/100 g DW). During the storage period, chlorogenic acid
levels were also increased for both coated and uncoating figs (from 1.21 to
3.14 mg/100 g DW and from 1.31 to 3.62 mg/100 g DW for uncoated and coated
figs, respectively), whereas gallic acid content was stable for coated figs. These
results imply that the modification of the internal atmosphere after applying
alginate-chitosan coating affects chlorogenic acid more than gallic acid.

2.3 Coumarins

Coumarins are characterized by a furan ring attached to carbon 6 and 7 (i.e., linear
type) or 7 and 8 (i.e., angular type) of a benzo-α-pyrone. Moraceae, Rutaceae,
Leguminosae, and Apiaceae are the four primary families of higher plants that con-
tain linear furanocoumarins, while the angular furanocoumarins are primarily found
in the Apiaceae and Leguminosae families. Psoralen, bergapten, xanthotoxin, and
isopimpinellin are the most abundant linear furanocoumarins existing in higher
plants (Hung et al., 2017). Coumarins are widely utilized for their antimicrobial and
anticancer activities (Hussain et al., 2019).
Psoralen and bergapten are the major coumarins detected in figs (Ficus carica).
In particular, fig leaves are the primary source of these compounds (Britto et al.,
2021; Chunyan et al., 2008; Kazemipoor et al., 2012; Teixeira et al., 2006; Wang
et al., 2017; Yu et al., 2020). According to Oliveira et al. (2012), the level of psoralen
and bergapten was relatively high in the fig leaves, corresponding to approximately
40–64% and 30–44% of total phenolic compounds, respectively. Besides fig leaves,
492 S. Kamiloglu and B. Akgun

fruits (Ammar et al., 2015; Qin et al., 2015), wood, and stem bark (Marrelli et al.,
2014; Rouaiguia-Bouakkaz et al., 2013) were also reported to contain psoralen and
bergapten. Psoralen content of fruit pulps was significantly higher than peels
(Oliveira et al., 2009). Similarly, both psoralen and bergapten levels were higher in
the wood than in the stem bark. Moreover, most dark-fruited fig trees produced
these two coumarins more than the green ones (Rouaiguia-Bouakkaz et al., 2013).
Besides psoralen and bergapten, Takahashi et al. (2014) identified a phenylpro-
panoid acid related to psoralen as psoralic acid glucoside. The authors suggested
that this compound could be a precursor of psoralen.
Psoralen possesses limited antioxidant activity due to its structure, which does
not contain any phenolic type of hydroxyl group that can act as a hydrogen donor
(Teixeira et al., 2009). Although coumarins have been described as low effective
antioxidants, these compounds were found to have an inhibitory effect on tumor
cells (Li et al., 2011).

3 Carotenoids

Carotenoids are fat-soluble pigments that provide the characteristic yellow, orange,
and red colors of many fruits, vegetables, and plant life. According to their chemical
composition, they are classified into two main groups: (i) xanthophylls and (ii) caro-
tenes. Lutein, α- and β-cryptoxanthin, zeaxanthin, canthaxanthin, and astaxanthin
are essential members of the xanthophylls group, whereas α- and β-carotene and
lycopene are the leading members of the carotene group (Kamiloglu et al., 2021).
As humans do not synthesize carotenoids, they need to be received through the diet
or via supplementation. The main potential benefits of carotenoids to human health
include contribution to (i) pro-vitamin A function, (ii) eye health, (iii) cognitive
performance, (iv) cardiovascular health, (v) bone health, (vi) sun protection, (vii)
weight management, (viii) immune function, (ix) cancer prevention and (x) infant
nutrition (Eggersdorfer & Wyss, 2018).
The carotenoids identified in figs include lutein, zeaxanthin, β-cryptoxanthin,
β-carotene, α-carotene, and lycopene (Table 21.2) (Amessis-Ouchemoukh et al.,
2017; Nawade et al., 2020; Petkova et al., 2019; Su et al., 2002; Yemiş et al., 2012).
Studies investigating the carotenoid content of figs revealed that the abundance of
these pigments depends on the variety. While the Australian figs contained lycopene
as the major carotenoid (0.32 mg/100 g) (Su et al., 2002), yellow Turkish figs (var.
Sarilop and Sarizeybek) (Yemiş et al., 2012) and green-purple figs from Israel (var.
Brown Türkiye) (Nawade et al., 2020) are reported to contain lutein as the most
abundant carotenoid (6.14–7.15 μg/g DW and 4.08–12.58 μg/g FW, respectively).
On the other hand, β-carotene was the major carotenoid in white Algerian figs.
(4.32 μg/100 g DW) (Amessis-Ouchemoukh et al., 2017). Overall, the total carot-
enoid content of fresh fig fruits (23.7–110 μg/g DW) was low compared to other
fruits (Veberic & Mikulic-Petkovsek, 2016).
21 Bioactive Compounds of Fig (Ficus carica) 493

Table 21.2 Carotenoid content of figs (Ficus carica L.)

Variety/Origin Skin color Form Compound(s) Content References
Unknown/Australia Unknown Fresh Lycopene 0.32 mg/100 g Su et al. (2002)
Lutein 0.08 mg/100 g
β-carotene 0.04 mg/100 g
α-carotene 0.02 mg/100 g
Cryptoxanthin 0.01 mg/100 g
Sarilop/Aydin, Yellow Fresh Lutein 6.14 μg/g DW Yemiş et al.
Türkiye Zeaxanthin 3.32 μg/g DW (2012)
β-cryptoxanthin 2.04 μg/g DW
β-carotene 1.40 μg/g DW
Dried Lutein 1.44–4.00 μg/g
DW
Zeaxanthin 0.76–2.00 μg/g
DW
β-cryptoxanthin 0.98–1.79 μg/g
DW
β-carotene 0.45–0.97 μg/g
DW
Sarizeybek/Aydin, Yellow Fresh Lutein 7.15 μg/g DW
Türkiye Zeaxanthin 2.20 μg/g DW
β-cryptoxanthin 1.85 μg/g DW
β-carotene 1.48 μg/g DW
Dried Lutein 1.85–5.25 μg/g
DW
Zeaxanthin 0.68–1.61 μg/g
DW
β-cryptoxanthin 1.00–1.78 μg/g
DW
β-carotene 0.47–1.22 μg/g
DW
Unknown/Beni White Fresh β-carotene 4.32 μg/100 g Amessis-
Maouche, Algeria DW Ouchemoukh
β-cryptoxanthin 2.14 μg/100 g et al. (2017)
DW
Croatian Yellow Yellow- Fresh Total 3.52 μg/g FW Petkova et al.
Giant/Plovdiv, green carotenoids (2019)
Bulgaria Frozen Total 1.73 μg/g FW
carotenoids
Jam Total 1.94 μg/g FW
carotenoids
Brown Türkiye/ Green- Fresh Total 6.64–18.39 μg/g Nawade et al.
Kibbutz, Machanaim, purple carotenoids FW (2020)
Israel (calculated as
the sum of
lutein,
β-carotene,
zeaxanthin, and
violaxanthin)
DW dry weight, FW fresh weight
494 S. Kamiloglu and B. Akgun

Besides the variety, several other factors affect the carotenoid content of fig
fruits. For instance, a study by Nawade et al. (2020) indicated that the accumulation
of carotenoid pigments decreases as the fruits mature. Accordingly, the total carot-
enoid contents of parthenocarpic and pollinated figs were reduced from 16.62–18.39
to 6.64–7.71 μg/g FW. Processing is another factor that affects the carotenoid con-
tent of fig fruits. Yemiş et al. (2012) monitored the changes in carotenoids during
sun-drying of figs and reported that the drying process reduced the total carotenoid
content of yellow Turkish figs by 78.5–81.5%. This reduction may be related to the
susceptibility of carotenoids to oxidative changes during drying due to their high
degree of unsaturation in chemical structure. In another study (Petkova et al., 2019),
the carotenoid content of frozen figs and fig jam was compared with fresh figs. The
results revealed that the carotenoid content in frozen fruits and jam was approxi-
mately 50% lower than that of fresh fruits. These results imply that both thermal and
cold processing decreases the carotenoid content of fig fruits.

4 Vitamins

Vitamins are substances essential for the development and growth of the body. They
are widely distributed in natural sources and have different chemical structures.
Vitamins are classified into two groups; water- or fat-soluble. Vitamins of A, D, E,
and K are fat-soluble. Vitamin C and the members of the vitamin B group, namely
thiamine (vitamin B1), riboflavin (vitamin B2), niacin (vitamin B3), pantothenic acid
(vitamin B5), vitamin B6, folate (vitamin B9), and vitamin B12 constitute the water-
soluble vitamins. The human body cannot synthesize most vitamins (except for
vitamin D and niacin), making them essential in the human diet. Vitamin D can be
synthesized in the body with adequate exposure to sunlight, and niacin synthesis is
related to a sufficient intake of tryptophan (Ball, 2005). Therefore, the optimum
values of daily vitamin intake can promote health and eliminate deficiency diseases.
Most foods can provide vitamins in variable quantities; nonetheless, there is no
single food containing all vitamins. Fruits generally provide a wide variety of vita-
mins such as vitamin A, vitamin B, vitamin C, and vitamin E. Hence, for disease
prevention, people should consume foods rich in vitamins (Uribe et al., 2017).
Nutritionally, fresh and dried figs are one of the richest plant sources in terms of
vitamins. For example, Arvaniti et al. (2019) stated the vitamin content of 100 g of
fresh figs as 142 IU of vitamin A, 2 mg of vitamin C, 0.060 mg of vitamin B1,
0.050 mg of vitamin B2, and vitamin content of 100 g dried figs as 10 IU vitamin A,
1.2 mg vitamin C, 0.085 mg vitamin B1, 0.082 mg vitamin B2..
According to Türkomp (2021), 100 grams of fresh Bursa black figs contain
2.1 mg vitamin C, 0.036 mg vitamin B1, 0.038 mg vitamin B2, 0.335 mg vitamin B3,
0.174 mg vitamin B6, 8 μg vitamin B9 and 11 retinol equivalents (RE) vitamin A. In
another database, it is shown that 100 grams of raw figs have 142 IU vitamin A,
2.0 mg vitamin C, 0.1 mg vitamin E, 4.7 mcg vitamin K, 0.1 mg vitamin B1, 0.1 mg
vitamin B2, 0.4 mg vitamin B3, 0.1 mg vitamin B6, 6 mcg vitamin B9 and 0.3 mg
21 Bioactive Compounds of Fig (Ficus carica) 495

Table 21.3 The vitamin content in 100 g raw figs (NutritionData, 2018a; Türkomp, 2021)
Vitamin Amount (TürKomp database) Amount (NutritionData)
Vitamin A 11 retinol equivalents (RE) 142 IU
Vitamin B1 0.036 mg 0.1 mg
Vitamin B2 0.038 mg 0.1 mg
Vitamin B3 0.335 mg 0.4 mg
Vitamin B5 – 0.3 mg
Vitamin B6 0.174 mg 0.1 mg
Vitamin B9 8 μg 6 mcg
Vitamin C 2.1 mg 2.0 mg
Vitamin E – 0.1 mg

vitamin B5 (NutritionData, 2018a) (Table 21.3). United States Department of

Agriculture reported 2 mg vitamin C, 7 mcg vitamin A, 4.7 mcg vitamin K, 6 mcg
vitamin B9 in 100 g raw figs, and 1.2 mg vitamin C, 0 mcg vitamin A, 15.6 mcg
vitamin K, 9 mcg vitamin B9 in 100 g dried figs (Caporuscio, 2019). Popova (2019)
determined vitamin C levels of commercially available fresh fruits as 1.1 mg/100 g
fresh sample and 1.9 mg/100 g fresh sample using two different assays. Similarly,
the vitamin C level of wild fig was stated as 3.3 mg/100 g of pulp by Kumari et al.
(2017). According to Morton (1987), 100 g of fresh fig fruit contained 20–270
I.U. vitamin A, 0.034–0.06 mg vitamin B1, 0.053–0.079 mg vitamin B2,
0.32–0.412 mg vitamin B3, 12.2–17.6 mg vitamin C. Guvenc et al. (2009) reported
that fig fruit contained some lipophilic vitamins (γ-tocopherol, δ-tocopherol, vita-
min D2, vitamin D3, α-tocopherol, α-tocopherol acetate, and vitamin K1) and vita-
min content varied in different sections. For example, vitamin K1 was the highest in
whole fig fruit, but it was relatively low in the skin. Also, the white part of the fig
contained the highest level of γ-tocopherol.
In the world, consumers have started to prefer dehydrated fruits. Suitable drying
techniques such as conventional hot air drying, sun drying, freeze-drying, and vac-
uum drying are commonly applied for obtaining dried figs. Dehydration removes
active water from fresh figs, which improves the product’s shelf life. However, high-
temperature drying for a long time may negatively affect the nutritional value of figs
(Abul-Fadl et al., 2015). This is because they were drying methods exposing the
food to air cause vitamin A and vitamin C losses due to oxidation. On the other
hand, Freeze-drying performed without oxygen does not result in vitamin C loss
(Ball, 2005). Chimi et al. (2005) stated that sun-drying could destroy the vitamins
of figs. Abul-Fadl et al. (2015) found that the L-ascorbic acid level of fresh figs
decreased significantly when different drying methods were applied. In this study,
L-ascorbic level was calculated as 11 mg/100 g dry weight basis of fresh fig, but
air-convection drying, vacuum-drying, microwave-air convection, and microwave-
vacuum drying reduced the amount of L-ascorbic acid by 90.2%, 76.6%, 63.1%,
and 52.1%, respectively. According to NutritionData (2018b), 100 g dried figs con-
tain 10.0 IU vitamin A, 1.2 mg vitamin C, 0.4 mg vitamin E, 15.6 mcg vitamin K,
0.1 mg vitamin B1, 0.1 mg riboflavin, 0.6 mg niacin, 0.1 mg vitamin B6, 9.0 mcg
496 S. Kamiloglu and B. Akgun

vitamin B9 and 0.4 mg vitamin B5, which proves that vitamin A and vitamin C level
are affected by the dehydration process. Similarly, Caliskan (2015) reported that
100 g of dried figs contained 0.68 mg of vitamin C. Also, Morton (1987) reported
80 I.U. vitamin A, 0.1 mg vitamin B1, 0.1 mg vitamin B2, 0.7 mg vitamin B3 and
0 mg vitamin C in 100 g dried figs.
By-products of figs (e.g., peel) can also be valuable in their vitamin contents.
Mahmoudi et al. (2018) investigated the vitamin C content of fig peels and pulps for
9 different cultivars. It was found that the cultivar of Onk Elhamam had the highest
vitamin C level as 10.67 ± 0.31 mg/100 fresh weight for peel and 9.33 mg/100 fresh
weight for pulp. On the other hand, the peel of the Hamra cultivar had the lowest
vitamin C level (1.33 mg/100 fresh weight).
In summary, vitamins are required nutrients for people, and fresh figs can be a
good source of vitamin A, B, C, and E. Also, dried figs and fig by-products contain
vitamins. However, dried figs have less vitamin A and C than fresh ones.

5 Organic Acids

Organic acids are acidic compounds significant for various industries such as food,
beverage, animal feed, and human health. These compounds impact the taste and
the microbiological stability of foods (Yang & Rainville, 2020). Organic acids may
occur naturally present in plant-based foods, or they can be formed during fermen-
tation. It is widespread to add these acids artificially to foods as acidulants, fla-
vorants, preservatives, or spoilage microorganisms’ growth inhibitors (Gurtler &
Mai, 2014). Some organic acids commonly used in foods are acetic, benzoic, citric,
formic, lactic, and propionic acid (Lianou et al., 2012). Organic acids can positively
affect people’s health. For instance, citric acid may prevent kidney stone formation.
However, some organic acids (e.g., oxalic acid) may lead to kidney stone formation
(Sukontaprapun et al., 2019). Malic and citric acid are the most widespread organic
acids in most fruits. However, other types of organic acids, such as tartaric acid, can
also be plentiful in some fruits. During the ripening of some fruits, the concentration
of malate and/or citrate decreases (Famiani et al., 2015).
Like other fruit species, figs contain organic acids impacting their quality. The
organic acid concentration of fruits is important since it affects the organoleptic
properties of fruits (Lee, 1993). Russo et al. (2010) stated that the organic acids
generally found in figs are tartaric, malic, citric, sorbic, and benzoic acids. Similarly,
Hussain and Othman (2011) determined the concentration of organic acids (citric
acid, fumaric acid, malic acid, and oxalic acid) in fig plants cultivated in orchard
adjacent to serpentine soil. The plant fruit‘s highest total organic acid content was
178 μg/g dry weight. Also, the lower amount of total organic acids was measured in
the branches with a 55.1 μg/g dry weight concentration. The contents of citric acid
varied significantly between fruits (28.7 μg/g) and leaves (14.3 μg/g). Non-
significant differences were detected between leaves (22.4 μg/g) and branches
(20.0 μg/g) in terms of fumaric acid. The highest oxalic acid concentration was
21 Bioactive Compounds of Fig (Ficus carica) 497

found in fruits (51 μg/g), and the branches (8.17 μg/g) contained the lowest levels
of oxalic acid.
Kelebek et al. (2018) detected two organic acids (citric and malic acid) in differ-
ent fig cultivars (Bursa Siyahi and Sarilop). In terms of organic acid profile, there
was no significant difference between fig varieties. However, considering the distri-
bution of organic acids, citric acid was dominant in fresh figs, while malic acid was
the most abundant in dried figs. The highest amount of citric acid was detected in
fresh Bursa siyahi figs. (4.99 g/kg), and dried Bursa siyahi figs contained the highest
concentration of malic acid (4.09 g/kg).
Slatnar et al. (2011) investigated the effect of the sun- or oven-drying of figs on
some organic acids (malic and citric acids) at different sampling dates. The level of
malic acid varied between 0.52 g/kg and 0.76 g/kg in fresh figs. Malic acid content
was found higher in dried figs, ranging between 3.11 g/kg and 9.07 g/kg. In fresh
figs, the citric acid content changed between 1.36 g/kg and 1.83 g/kg. Similarly, the
citric acid level increased in dried figs (changing between 3.33 g/kg and 10.5 g/kg).
Shiraishi et al. (1996) elucidated organic acid profiles in the juice of fig fruits
from Japan (Brown Türkiye and Houraishi cultivars). In the juice of all figs, variet-
ies studied, acetic, butyric, oxalic, malonic, succinic, fumaric, malic, and citric
acids were detected. No propionic and lactic acids were found, but glyoxylic and
tartaric acid was only determined in a few examined cultivars. In another study by
Trad et al. (2014), organic acid contents of figs from Tunisia were determined. The
concentrations of organic acids were relatively higher in the Zidi variety (0.43%
citric acid and 0.17% malic acid). On the other hand, the lowest citric acid (0.23%)
and malic acid (0.11%) were detected in the Bouhouli and Bidhi varieties.
Moreover, Melgarejo et al. (2001) investigated organic acids of first and second
crop fig juices from 4 different cultivars (Colar, Florancha, Tio Antonio, and
Gobernador). Acetic and fumaric acid were not generally detected in fig juices. On
the other hand, malic (0.0093–0.0318%), oxalic (0.0036–0.0398%), citric
(0.0056–0.0399%), and tartaric (0.0091–0.0346%) acids were reported at varying
levels in analyzed fig juices.
Mohammed et al. (2020) identified organic acid levels of fig organs (leaf, stem,
and fruit) using HPLC. Organic acids are distributed unequally in different parts of
fig organs. Malic (61.4 μg/g) and oxalic (51.0 μg/g) acids were dominant organic
acids in the studied fig fruits. Citric acid content showed a difference in different fig
parts. The highest level of citric acid was found in fig fruit (28.7 μg/g), and the low-
est level of citric acid was detected in the leaf (14.3 μg/g). The highest fumaric acid
concentration was obtained in the fruit (36.9 μg/g). The total organic acid content
was determined in fruits as 178 μg/g, leaves as 71.7 μg/g, and stems as 55.1 μg/g.
Shin et al. (2015) investigated changes in organic acids in 120 figs at six different
maturation stages. In the final stage, figs mainly contained citric acid, malic acid,
and tartaric acid. In another study, Zhang et al. (2020) determined organic acid com-
ponents in figs at two ripening stages using HPLC system. The organic acids,
namely tartaric acid, quinic acid, malic acid, citric acid, fumaric acid, and succinic
acid, were detected in the peel and pulp of Sichuan fig from China. The malic acid,
citric acid, and quinic acid ranges were 104.8–625.6 mg/100 g fresh weight,
498 S. Kamiloglu and B. Akgun

9.78–555 mg/100 g fresh weight, and 104.3–311.1 mg/100 g fresh weight in the
pulp, respectively. The range of fumaric (0.38–2.92 mg/100 g fresh weight) and
tartaric acid (6.54–83.2 mg/100 g fresh weight) in the peel were considerably lower
than other organic acids.
Overall, fig contains organic acids, which are significant for human health.
Organic acids such as citric, malic, fumaric, tartaric, and oxalic acid are commonly
found in different parts of fig organs. Different types of factors such as ripening and
drying may affect the organic acid content of figs.

6 Terpenes

Terpenes are hydrocarbons with the formula (C5H8)n (Breitmaier, 2006), and accord-
ing to their carbon units, they are classified as hemiterpenes (C5), monoterpenes
(C10), sesquiterpenes (C15), diterpenes (C20), sesterterpenes (C25), triterpenes
(C30) and carotenoids (C40) (Ullah & Khan, 2020). Terpenes (also known as terpe-
noids) are naturally occurring compounds that are the primary components of
essential oils, and they play a role in the fragrance, taste, and pigments of plants
(Cox-Georgian et al., 2019). Plant oil containing terpenes has traditionally been
used to treat various diseases (Cho et al., 2017). Many terpenes are bioactive
(Kupska et al., 2016), and figs are a significant source of various bioactive com-
pounds, including terpenes (Solana & Romano, 2019). Studies on the aroma profile
of figs showed that certain compounds such as aldehydes, alcohols, ketones, and
terpenes constitute fig odor. Volatile compound quantities may change in different
fig cultivars, fresh and dried figs (Palassarou et al., 2017).
Zidi et al. (2021) investigated the volatile organic compound (VOC) profiles of
figs from 3 cultivars (Taamriwthe, Azegzaw, and Averkane) at 3 ripening stages
(unripe, ripe, and fully ripe). Figs were harvested from Algeria in 2018. Six terpene
compounds (limonene, linalool, epoxylinalol, α-Santalene, 1,8-cineol, and
β-caryophyllene) were examined in volatile fig compounds. The most abundant ter-
pene in the Azegzaw fig was linalool, and only 2 terpenes (linalool and 1,8-cineol)
were detected in the odor of Averkene fig. Moreover, β-caryophyllene was only
identified in unripe and ripe Taamriwthe fig, and α-Santalene was only found in ripe
Taamriwthe fig.
Russo et al. (2017) characterized volatile compounds of white figs cv. “Dottato”
from Southern Italy and commercial dried figs from Türkiye. In addition to alde-
hydes, alcohols, and ketones, terpenes were found as another class responsible for
fig aroma. D-limonene, alpha-pinene, and 3-carene were identified in Dottato figs
dried by the sun, Dottato figs oven-treated after the sun-drying, and Turkish sam-
ples. D-limonene was detected in all samples, and it was the most plentiful (3.64 mg/
kg) in figs oven-treated after the sun-drying. Alpha-pinene (0.42 mg/kg) and
3-carene (0.17 mg/kg) were only detected in Turkish fig samples. It was noticed that
thermal treatment led to a significant increase in terpene content.
21 Bioactive Compounds of Fig (Ficus carica) 499

Pereira et al. (2020) determined volatile compounds of fresh fig samples from 9
cultivars: early cultivars (San Antonio, Blanca Bétera, Brown Türkiye, Tres Voltas
L’Any), mid cultivar (Banane), and late cultivars (Cuello Dama Blanco, Cuello
Dama Negro, Colar Elche, and De Rey). They detected monoterpenes (unknown
monoterpene (min-max: 0–0.39%), α-pinene (min-max: 0–0.65%), limonene (min-
max: 0–1.18%) and linalool (min-max: 0–4.35%)). They found that terpenes were
the best physicochemical property to assess the overall acceptability of figs, which
was highly related to sensory attributes of flesh color and fruit flavor.
Gozlekci et al. (2011) investigated the aroma profile of peels and pulps of 4 fig
cultivars (“Bursa Siyahi”, “Karabakunya”, “Sari Lop” and “Sultan Selim”) from
Türkiye by using Headspace Gas Chromatography-Mass Spectrophotometer (HS/
GC-MS) technique. Among terpenes, β-caryophyllene and germacrene D were the
main volatile compounds found in peel and pulp fig. The peel of “Karabakunya”
contained the highest amount of β-caryophyllene (48.6%), and the peel of “Sari
Lop” had the highest germacrene D composition (9.19%). Limonene content
changed between not detected and 0.59% in a different part of figs. Total terpenes
(60.57%) were the highest in the peel of figs from the Sultan Selim cultivar.
Mujić et al. (2012) analyzed the aroma profile of fresh figs (cultivar Petrovaca
Crna) from Croatia, fresh figs frozen in liquid nitrogen, and dried figs (dark variety
Petrovaca Crna) by HS-SPME followed by GC-MS. In tested fig samples, 80 volatile
compounds were identified from different chemical groups (alcohols, aldehydes,
esters, acids, terpenes, terpenic compounds, and other compounds). Linalool
(3,7-dimethyl-1,6-octadien-3-ol) was the most abundant terpenic compound. They
also found that the peak area of terpenes and terpenic compounds declined with
dehydration.
The aroma profile of fresh fruits (pulp and peel) and leaves of Portuguese Ficus
carica L. white (“Pingo de Mel” and “Branca Tradicional”) and dark (“Borrasota
Tradicional”, “Verbera Preta” and “Preta Tradicional”) varieties were determined by
Oliveira et al. (2010a, b). 59 aroma compounds from different chemical classes
(aldehydes, alcohols, ketones, esters, monoterpenes, sesquiterpenes, and noriso-
prenoids) were found by HS-SPME/GC-IT-MS in 5 Portuguese varieties of F. carica
leaves and fruits. 2 monoterpenes named limonene and menthol and 17 sesquiter-
penes such as copaene, germacrene D, (+)-Ledene and alloaromadendrene were
reported in five Portuguese varieties of F. carica leaves. Also, 7 monoterpenes and 7
sesquiterpenes were found in five Portuguese varieties of F. carica fruits (peels and
pulps). Furthermore, Trad et al. (2012) identified volatile compounds in pollinated
and non-pollinated Tunisian figs from 3 cultivars (Bouhouli, Zidi, and Thgagli).
γ-Sesquiterpene was only found in pollinated (1.27 μg/kg) and non-pollinated (2 μg/
kg) figs from Bouhouli cultivar. Germacrene D (80 μg/kg) and linalool (8 μg/kg)
found a higher amount in pollinated Bouhouli fig than in other samples.
Song et al. (2001) determined volatiles of figs (receptive, postpollinated, and
postparasitized) and leaves of Ficus hispida. Linalool was the main component of
steam-distilled oil of either male or female receptive figs. The volatile composition
of figs changed from a receptive to a postpollinated state. It was thought that lin-
alool, linalool oxide, alpha-terpeneol, and 2,6-dimethyl-1,7-octadiene-3,6-diol
acted as the attractants of the wasps.
500 S. Kamiloglu and B. Akgun

Yao et al. (2021) identified aroma compounds of Xinjiang dried figs by GC-MS
and GC-O. Myrcene, styrene, α-copaene, β-caryophyllene, and δ-cadinene were
reported under the terpenes and terpenic compounds studied in the figs. Moreover,
Grison-Pigé et al. (2002) identified the blends of volatile compounds emitted by
receptive figs of 20 Ficus species. Different terpenes (acyclic monoterpenes, cyclic
monoterpenes, and sesquiterpenes) were detected. These studies showed that, figs
may contain terpenes such as limonene, linalool, β-caryophyllene, and Germacrene D.

7 Phytosterols

Phytosterols comprise both plant sterols and plant stanols. Although the human
body cannot synthesize them, they can be obtained from plant-based foods such as
vegetables, fruits, nuts, and cereals (Martianto et al., 2021; Ras, 2014). In a typical
western diet, the daily consumption of plant sterols is 200–400 mg. In the diet, the
most widespread sterols are β-sitasterol, campesterol, and stigmasterol (Gupta et al.,
2011). In addition, phytosterols offer different health benefits, such as lowering the
serum cholesterol effect and preventing colon cancer (Han et al., 2008). Thus, phy-
tosterols should be added to the diet.
In fruits, the total plant sterol contents generally varied from 1.6 to 32.6 mg/100 g
(Han et al., 2008). Figs are good sources of phytosterols (Kumari et al., 2018), and
different studies have been performed to identify the phytosterols in the figs and fig
leaves. Gupta et al. (2011) stated total phytosterols in figs as 22 mg/100 g. Jeong and
Lachance (2001) determined the sterol profiles of various parts of fig tree branches
(bark, stem, and pith). It was found that all parts contained campesterol, stigmas-
terol, and sitosterol, which are the most common phytosterols in nature. β-Amyrin
and psi-taraxasterol are also found in all parts. Moreover, Oliveira et al. (2010a)
determined 7 phytosterols (betulol, lupeol, lanosterol, lupeol acetate, β-amyrin,
β-sisterol, and α-amyrin) in a representative sample from F. carica latex by gas
chromatography-ion trap mass spectrometry (GC-ITMS).

8 Fatty Acids

Fatty acids, an important component of lipids, can be categorized as saturated,

monounsaturated, polyunsaturated, and trans fats. Foods contain over 20 types of
fatty acids. Palmitic acid (saturated), stearic acid (saturated), oleic acid (monoun-
saturated), linoleic acid (polyunsaturated), linolenic acid (polyunsaturated), arachi-
donic acid (polyunsaturated), eicosapentaenoic acid (polyunsaturated) and
docosahexaenoic acid (polyunsaturated) are most well-known fatty acids (White,
2009). Omega-3 (e.g. α-linolenic acid (18:3n-3), stearidonic acid (18:4n-3), eicosa-
pentaenoic acid (EPA; 20:5n-3) and docosahexaenoic acid (DHA; 22:6n-3)) (Calder,
2013) and omega-6 (e.g. linoleic acid (LA; 18:2n-6), gamma-linolenic (18:3n-6)
21 Bioactive Compounds of Fig (Ficus carica) 501

and arachidonic acid (ARA; 20:4n-6) are the essential fatty acids (Innes & Calder,
2018). They cannot be synthesized in the body; therefore, people must get these
polyunsaturated fatty acids from foods. Monounsaturated fatty acids and polyun-
saturated fatty acids serve many health benefits, such as decreasing coronary heart
disease risk; thus, they are necessary for the human diet (White, 2009). There is an
increase in saturated fat, omega-6 fatty acids, and trans fatty acids intake in indus-
trialized societies and a decrease in omega-3, fruit and vegetable intake. This situa-
tion leads to an imbalance in the ratio of omega 6 to omega 3 (Simopoulos, 2009),
which shows the necessity of changing dietary trends and food consumption habits.
For centuries, fig fruit has been an indispensable part of the health-promoting
Mediterranean diet. The fruits, roots, and leaves of fig trees have a number of thera-
peutic properties and they are used to treat cardiovascular, respiratory, and gastroin-
testinal disorders (Nakilcioğlu-Taş, 2019). In addition, fruits are rich in nutrients
such as vitamins, phenolic acids, and fatty acids (Icyer et al., 2017). Awareness of
the importance of fatty acids in human health has increased recently (Kaur et al.,
2014), and therefore several studies have been conducted to investigate the fatty
acid content of figs.
Guvenc et al. (2009) reported that the fig fruits from Adiyaman district, Türkiye
were rich in fatty acids. Gas chromatographic (GC) analysis of fatty acid methyl
esters was used, and both non-essential (palmitic, stearic, and oleic acid) and essen-
tial (linoleic and linolenic acid) fatty acids were identified in fig fruit. Linoleic
(18:2n-6), linolenic (18:3n-3), oleic (18:1n-9), and palmitic (16:0) acids were found
at high levels in all fig parts (whole fig fruit, skin, white part, and red part). Linoleic
acid levels changed between 19.35% and 24.38% in different fig parts. The highest
linolenic acid content (39.3%) was detected in the red part of the fig fruit.
In another study, Palmeira et al. (2019) assessed the fatty acid content of parts of
a Portuguese common fig variety by GC coupled to a flame ionization detector
(FID). They detected 23 fatty acids in samples. There were differences in the fatty
acid composition of peel and pulp extracts. The α-linolenic acid (18:3n3) was found
to be the most abundant fatty acid in fig peel (28.0%) and pulp (51.8%), followed by
linoleic (18:2n6), palmitic (16:0), and oleic (18:1n9) acids.
According to Solana and Romano (2019), polyunsaturated fatty acids constituted
84% and 69% of total fatty acids in dried and fresh fi. Oleic acid was the most plen-
tiful in fig fruits concerning monounsaturated fatty acids. Moreover, Jeong and
Lachance (2001) determined fatty acids in dried fig fruit as their methyl esters using
the GC system. Myristic (14:0), palmitic (16:0), stearic (18:0), oleic (18:1), linoleic
(18:2) and linolenic (18:3) acids formed its fatty acid composition. The most domi-
nant fatty acid was linolenic acid (53.1%), followed by linoleic acid (21.1%), pal-
mitic acid (13.8%), and oleic acid (9.8%). It was found that 84% of the total fatty
acids were unsaturated fatty acids.
ANSES-CIQUAL food composition table showed that 100 g of dried figs con-
tained 0.65% myristic acid, 12% palmitic, 3% stearic acid, 18% oleic acid, 22%
linoleic acid, and 2% linolenic acid (Anonymous, 2021). Khadhraoui et al. (2019)
found that fatty acid contents of dry fig cultivars were composed of saturated fatty
acids, monounsaturated fatty acids, and polyunsaturated fatty acids. They identified
502 S. Kamiloglu and B. Akgun

palmitic acid (16:0), oleic acid (18:1), linoleic acid (18:2), linolenic acid (18:3), and
arachidic acid (20:0) in 9 sun-dried fig cultivars by GC-MS. On average,
0.35 ± 0.02 g/100 g dry matter (DM) linoleic acid and 0.39 ± 0.04 g/100 g DM lino-
lenic acid were found, which were the dominant fatty acids in all samples. Arachidic
acid (C:20) was detected in all fig samples at a low level.
Depending on the maturity and size of a fruit, a fig may contain a large number
of small seeds (Kolesnik et al., 1986). For example, Joseph and Raj (2011) stated
that dried seeds have a substantial amount of oil containing linoleic (33.7%), linole-
nic (32.9%), oleic (18.9%), palmitic (5.23%), stearic (2.18%) and arachidic (1.05%)
acid. Likewise, Icyer et al. (2017) found linolenic (41.1%), linoleic (30.9%), oleic
(30.9%), and palmitic (7.16%), and stearic (2.9%) and no trans fatty acid in fig
seed oil.
Güven et al. (2019) analyzed the fatty acid content of fig seed oil from Türkiye
using GC. Nine major fatty acids (palmitic acid, stearic acid, oleic acid, linoleic
acid, α-linolenic acid, γ-linolenic acid, arachidic acid, eichosenoic acid, and behenic
acid) were identified in fig seed oil. Alpha-linolenic acid (18:3n-3) (32–50%) and
linoleic acid (18:2n-6) (20–35%) were the dominant fatty acids. Moreover,
Nakilcioğlu-Taş (2019) detected 14 fatty acids in fig seed oil. Fig seeds from a dif-
ferent location (Incirliova, Germencik, and Nazilli) could be a good source of
γ-linolenic acid (18:3n-6), ranging from 37.8% to 41.8%. Also, the level of linoleic
acid (18:2n-6) was the second-highest varying between 31.8% and 37.9%. On the
other hand, the lowest amount (0.02–0.03%) was reported for arachidic acid.
Ergun and Bozkurt (2020) investigated the fatty acid composition of fig seed oil
with GC and detected 19 different fatty acids in fig seed oil. The most abundant fatty
acid was α-linolenic acid (26.3% of total fatty acids). Linoleic acid (18:2n-6)
(24.2%) and oleic acid (18:1n-9) (19.6%) were the other dominant fatty acids in fig
seed oil. Furthermore, Hssaini et al. (2020b) analyzed the fatty acid content of seeds
oil of four fig cultivars. Eleven fatty acids were identified, and linolenic acid was the
most dominant fatty acid varying between 38.4% and 43.5% in all samples. The
average contents of linoleic acid were in the range of 28.9–34.5%. The third impor-
tant fatty acid was oleic acid ranging from 13.4% to 15.6%.
Uysal et al. (2015) studied the fatty acid composition of fig tree leaves collected
from the Mediterranean Region of Türkiye. It was reported that the predominant
fatty acid in fig leaves was α-linolenic acid (18:3n-3) (25.6%), followed by palmitic
acid (16:0) (25.2%), γ-linolenic acid (18:3n-6) (13.7%), linoleic acid (18:2n-6)
(10.6%), stearic acid (18:0) (8.35%) and oleic acid (18:1n-9) (4.19%).
Studies demonstrated that different parts of fig fruits and fig by-products (e.g.,
seeds, leaves, and peel) could be important sources of some fatty acids. Generally,
α-linolenic acid, γ-linolenic, linoleic acid from polyunsaturated fatty acids, oleic
acid from monounsaturated fatty acids, and palmitic acid from saturated fatty acids
are found in high quantity in figs and fig by-products. Thus, the consumption and
use of these products could be good dietary essential fatty acid sources for humans.
21 Bioactive Compounds of Fig (Ficus carica) 503

9 Conclusions

Fig (Ficus carica) is one of the most abundant fruits in the Mediterranean diet that
promote human health owing to its bioactive compounds such as polyphenols,
carotenoids, vitamins, organic acids, triterpenoids, phytosterols, and fatty acids.
Flavonoids, including anthocyanins, flavonols, flavanols and flavones, and phenolic
acids and coumarins, are the main polyphenolic compounds present in figs. The
levels of bioactive compounds in figs depend on the variety, cultivation, environ-
mental conditions, and processing parameters. Dark-colored fruits contain higher
bioactive compound concentrations than figs with light-colored skin. Therefore, the
skins of fig fruit should not be discarded as they contain several health-promoting
nutrients. Accumulation of polyphenols in fig fruits increases during fruit ripening,
whereas the levels of carotenoids reduce as the fruits mature. Drying process results
in the reduction of some bioactive compounds, especially anthocyanins and carot-
enoids, as well as vitamins. On the other hand, the content of specific bioactive
compounds such as organic acids is reported to enhance in dried figs. Overall, both
fresh and dried figs are good sources of bioactive compounds that possess health-
promoting properties.

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Chapter 22
Fig Volatiles

Mustafa Kiralan, Sündüz Sezer Kiralan, and Onur Ketenoglu

1 Introduction

Fig (Ficus carica L.) is a member of the botanical family Moraceae and is grown in
many parts of the world with moderate climates. Fig has worldwide importance for
dry and fresh consumption (Dueñas et al., 2008), and the consumers prefer the fresh
form. According to the reported data by FAO in 2019 (FAO, 2021), the worldwide
fig production is 1.315.588 tonnes. Turkey is a major producer of edible figs with
310,000 tonnes, followed by Egypt and Morocco (the production quantity of these
countries are 225,295 tonnes and 153,472 tonnes, respectively).
The different parts of figs have many positive effects on human health, such as
anti-inflammatory, antipyretic, purgative, aphrodisiac, anti-cancer, antidiabetic,
hepatoprotective, and anti-angiogenic properties (Ghanbari et al., 2019; Patil &
Patil, 2011; Rahmani & Aldebasi, 2017). Figs are an excellent source of nutrition
due to their rich composition in minerals, vitamins, and dietary fiber (Badgujar
et al., 2014; Crisosto et al., 2011). Carbohydrates constitute a large part (73.5%) of
the dried figs, while protein content is reported to be 4.67%, and the dietary fiber
content of dried figs is 3.68% (Soni et al., 2014). Fructose and glucose are the most
common sugars in fig fruit (Slatnar et al., 2011). Çalişkan and Aytekin Polat (2011)
emphasized that major sugars were fructose (∼56%) and glucose (∼43%) in
seventy-six Turkish fig accessions. The highest values in fructose and glucose were
11.9 g/100 g and 9.7 g/100 g, respectively.

M. Kiralan (*) · S. S. Kiralan

Department of Food Engineering, Balikesir University, Faculty of Engineering,
Balikesir, Turkey
O. Ketenoglu
Faculty of Agriculture, Department of Food Engineering, Eskisehir Osmangazi University,
Eskisehir, Turkey

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 513
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_22
514 M. Kiralan et al.

Besides these nutritional compounds, fig; contains different bioactive com-

pounds such as phenolic compounds, phytosterols, volatiles, organic acids, hydro-
carbons, and aliphatic alcohols. The presence and quantity of these phytochemicals
depend on the different parts of the fig. Most of these compounds are present in
latex, followed by leaves, fruit, and root (Badgujar et al., 2014; Mawa et al., 2013).
The content of total phenolics, total anthocyanins, and total flavonoids in Tunisian
fig ecotypes varied from 51.5 to 100 mg gallic acid equivalents (GAE) on a fresh
weight (FW) basis, 2.57 to 11.6 mg of cyanidin-3-glucoside equivalents (CGE) per
g FW, 0.33 to 17.59 mg of quercetin equivalent in mg per g FW, respectively (Aljane
et al., 2020).
Among phenolic acids, chlorogenic acid was the major phenolic acid in three fig
cultivars grown in Slovenia, followed by gallic acid (Veberic et al., 2008). As for
organic acids, malic acid was the main compound in fig samples, representing
24.7–58.7% of the total organic acids content (Slatnar et al., 2011). Cyanidin-3-
glucoside, cyanidin-3,5-diglucoside, cyanidin-3-rutinoside (major anthocyanin),
and pelargonidin-3-glucoside were identified in three fresh common fig cultivars
grown in Turkey, additionally; lutein, zeaxanthin, β-cryptoxanthin, and β-carotene
were the identified carotenoids in yellow fig cultivars grown in Turkey (Yemiş
et al., 2012).
Latex is a part of fresh fig, consumed in small amounts with fresh fig. Phytosterols,
especially β-sitosterol and lupeol, are the major sterols present in the latex of fresh
figs. Cysteine and tyrosine are present as the major amino acids among 13 amino
acids identified in F. carica latex. Besides, F. carica latex was reported to contain
saturated fatty acids such as palmitic, arachidic, and behenic acids (Oliveira,
et al., 2010a).
Alongside nutrients and functional compounds, volatile compounds are impor-
tant constituents that contribute to the unique aroma of fig. This section will discuss
the chemical composition of volatiles, extraction methods, and usage of fig volatiles
in different industries.

2 Chemical Composition of Volatiles

Volatile compounds contribute to the characteristic aroma and flavor of fruits.

Numerous chemical compounds such as esters, alcohols, aldehydes, ketones, lac-
tones, terpenoids, and apocarotenoids constitute the aroma profile of fruits (El Hadi
et al., 2013). Volatile compounds have been used to determine fruit quality.
Especially, individual volatile compounds could be monitored and evaluated in
fruits (Plutowska & Wardencki, 2007). Volatile compounds are also an important
part of fig composition. The chemical composition of fig volatiles changes with
climatological conditions, cultivar, maturity, processing, and storage conditions
(Grison-Pigé et al., 2002; Mujić et al., 2012, 2014).
Gozlekci et al. (2011) reported that aldehydes and terpenes were the major
chemical groups responsible for the aroma of peels and pulps in fig fruits from four
22 Fig Volatiles 515

CH3

H2C O
H H
CH3 H
CH3

b-caryophyllene E-2-hexenal Germacrene D t-elemene

CH3

Limonene Menthol

Fig. 22.1 Chemical structures of important volatiles in different parts of figs

commercial fig cultivars (Bursa Siyahi, Karabakunya, Sari Lop, and Sultan Selim)
growing in Turkey. According to the researchers, total terpenes were higher in the
peel than in the pulp. β-Caryophyllene was the major terpene in all samples, as also
found in high amounts in the peels of figs (up to 48.65% of total volatiles). E-2-
Hexenal was found as the major aldehyde, and its level increased up to 20.0%.
Oliveira et al. (2010c) identified 59 volatile and semi-volatile compounds in the
leaves and fruits (peels and pulps) from Portuguese fig cultivars. The number of
identified compounds was found to be higher in leaves (40 different compounds)
than in pulps (30) and peels (27). Aldehydes and monoterpenes were abundantly
present in pulp and peels, while these compounds were present at lower leaf concen-
trations. Sesquiterpenes were the major class of compounds in the leaves.
Germacrene D, β-caryophyllene, and τ-elemene (Fig. 22.1) were the predominant
sesquiterpenes in the leaves of all cultivars. On the other hand, monoterpenes were
found at higher concentrations in pulps and peels of figs. The major monoterpenes
present in pulps and peels were limonene and menthol.

3 Extraction Techniques

Different extraction techniques influence volatile compounds in figs. However,

most conventional methods require large volumes of organic solvents and result in
significant waste, requiring longer extraction time and trace levels of various com-
pounds. From these aspects, solid-phase microextraction (SPME) is a solvent-free,
cost-effective, rapid, sensitive technique (Jalili et al., 2020; Kumar et al., 2008).
516 M. Kiralan et al.

Table 22.1 Composition of fig volatiles depends on different parts of the fruit and different
extraction techniques
Part of
figs Extraction method Major volatiles Reference
Leaves Hydrodistillation (Z)-3-Hexenyl benzoate (19.8%), n-Tetracosane Ayoub
(11.6%), n-Hexadecanoic acid (9.2%), (E)-2- et al.
Hexenal (7.2%) (2010)
Leaves Hydrodistillation Ficusin (32.8%), Caryophyllene oxide (9.0%), Nafis et al.
Bergapten (6.1%) (2019)
Fruits Hydrodistillation Fruits Li et al.
Leaves Furfural (10.55%), 5-methyl-2-furaldehyde (10.1%), (2012)
and benzeneacetaldehyde (6.59%)
Leaves
Psoralen (10.12%), β-damascenone (10.17%),
benzyl alcohol
(4.56%), behenic acid (4.79%), and bergapten
(1.99%)
Fruits Hydrodistillation Fruits Soltana
Leaves Cedrol (43.8%), α-terpinyl acetate (22.5%), manoyl et al.
oxide (12.9%), α-pinene (9.3%), abietadiene (2017)
(8.1%),trans-calamenene (3.9%) and nheneicosane
(3.5%)
Leaves
Cedrol (38.9%), manoyl oxide (24.8%), α-terpineol
acetate (21.7%), abietatriene (11.8%), γ-muurolene
(7.4%), α-pinene (6.1%), pentadecanal (5.2%) and
nonadecanal (2.3%).
Latex Headspace (1) sesquiterpenes (ca. 91% of identified Oliveira,
Solid-Phase compounds) et al.
Microextraction germacrene D (2010b)
(HS-SPME) cadinene
τ-muurolene
(2) alcohols (ca. 4%)
1-hexanol
1-heptanol
(3) Ketones represented the minor components
(<0.1%)
6-Methyl-5-hepten-2-one
Pulp Headspace β-Caryophyllene (15.9–37.4%), (E)-2-Hexanal Gozlekci
Solid-Phase (11.0–20.0%), 2-Furancarboxaldehyde, et al.
Microextraction 5(hydroxymethyl), (5.96–13.5%), benzaldehyde (2011)
(HS-SPME) (5.05–8.45%), furfural (0.30–3.19%)
Peel Headspace β-Caryophyllene (21.1–48.6%), Gozlekci
Solid-Phase 2-Furancarboxaldehyde, 5(hydroxymethyl) (not et al.
Microextraction detected-4.56%), furfural (2.56–3.65%), (E)-2- (2011)
(HS-SPME) Hexanal (not detected-1.77%), benzaldehyde (not
detected-0.24%)
(continued)
22 Fig Volatiles 517

Table 22.1 (continued)

Part of
figs Extraction method Major volatiles Reference
Whole Headspace (1) Ester Pereira
Solid-Phase ethyl acetate (0.67–65.26%) et al.
Microextraction (2) Furans (2020)
(HS-SPME) 5-Hydroxymethylfurfural (0–33.38%)
(3) Pyranones
3-Hydroxy-2,3-dihydromaltol (0.36–25.06%)
(4) Aldehydes
Benzaldehyde (1.45–32.44%)
Flesh Headspace Cuello Dama Negro San Antonio Villalobos
Solid-Phase Hydrocarbons (2.10%) Hydrocarbons et al.
Microextraction Hexane (0.90%) (17.92%) (2018)
(HS-SPME) Aldehydes (12.15%) Hexane (12.16%)
3-Methyl butanal (0.06%) Aldehydes
Hexanal (7.64%) (17.88%)
Esters (1.14%) 3-Methyl butanal
Ethyl acetate (0.59%) (11.18%)
Pyranones (70.88%) Hexanal (1.36%)
3-Hydroxy-2,3- Esters (43.96%)
dihydromaltol Ethyl acetate
(54.24%)-Maltol (0.52%) (43.15%)
Pyranones (2.08%)
3-Hydroxy-2,3-
dihydromaltol
(0.02%)
Maltol (2.01%)

Both traditional and modern extraction techniques have been used to evaluate the
chemical composition of figs. Table 22.1 summarizes the volatile compounds
depending on the part of the fruit and extraction technique. For example, Li et al.
(2012) emphasized that furfural and 5-methyl-2-furaldehyde were the major vola-
tiles in the fruit part, while psoralen and β-damascenone were predominant volatiles
in the leaves of figs. In another study in which hydrodistillation was used, the fruits
were rich in cedrol (43.8%), α-terpinyl acetate (22.5%), and manoyl oxide (12.9%);
likewise, cedrol (38.9%), manoyl oxide (24.8%), α-terpineol acetate (21.7%) were
found as major compounds in the leaves of figs (Soltana et al., 2017).
Headspace solid-phase microextraction (HS-SPME) significantly improves the
determination sensitivity of volatile compounds in figs. Gozlekci et al. (2011) stated
that a bicyclic sesquiterpene, namely β-caryophyllene, was present in high percent-
ages of pulp and peel parts of figs. (15.9–37.4%, 21.1–48.6%, respectively).
β-Caryophyllene is reported to contribute a unique aroma to the essential oils of
most plants (Scandiffio et al., 2020). The wooden odor is a characteristic of
β-caryophyllene, making this volatile preferable in cosmetic and food additives.
Besides, this substance is approved as a flavoring agent by official authorities such
as the Food and Drug Administration (FDA) and the European Food Safety Authority
(EFSA). β-caryophyllene is also reported to have such biological activities as
518 M. Kiralan et al.

anti-inflammatory, anticarcinogenic, antimicrobial, antioxidant, antihyperglycemic

effects, and analgesic activities (Fidyt et al., 2016; Hashiesh et al., 2020; Moo et al.
2020; Scandiffio et al., 2020).
Villalobos et al. (2018) revealed that esters were the most abundant chemical
group in the flesh of figs from the San Antonio cultivar, while the major chemical
group was pyranones in the Cuello Dama Negro cultivar using the HS-SPME
method. The most abundant compound was 3-hydroxy-2,3-dihydromaltol, at
54.24% of the total area in the Cuello Dama Negro cultivar, while ethyl acetate
(43.15%) was the predominant volatile compound present in the San Antonio
cultivar.
Latex is widely distributed and produced from the fig tree and consists of second-
ary metabolites such as terpenoids, phenolics, and essential fatty acids (Lazreg Aref
et al., 2011). Oliveira et al. (2010b) used the HS-SPME method to identify volatiles
in fig latex. The major chemical group was sesquiterpenes which comprised 91% of
the identified compounds. According to these authors, germacrene D, cadinene, and
τ-muurolene were the major sesquiterpenes.

4 Factors Affecting Fig Volatiles

Different factors such as processing conditions influence the volatiles of figs. Mujić
et al. (2014) examined the effect of pre-treatments such as sulfur dioxide, citric acid,
and ascorbic acid on the volatiles of figs before drying. Benzaldehyde was found as
the major volatile in all treated and untreated dried figs (56–82.7 mg/kg), followed
by hexanal (7.20–55.7 mg/kg). Although the pre-treatments provided some advan-
tages, such as decreasing the drying time, they caused a decrease in the aldehydes
part of the aroma components. On the other hand, ascorbic acid application pre-
served esters which are responsible for the desired aroma in figs from Bružetka
Bijela variety, whereas, in the case of Zimnica, sulfur dioxide showed a protective
effect on esters compared to ascorbic acid (Mujić et al., 2014).
Russo et al. (2017) evaluated the effect of drying methods on the volatiles of figs.
Aldehydes were found as the major chemical group in sun-dried (approximately
70% of the total volatiles) and oven-treated samples after sun drying (approximately
80% of the total volatiles), especially hexanal and benzaldehyde were the most
abundant volatiles. A significant increase was observed in the samples treated with
the oven after sun-drying, while a decrease was observed in alcohols and esters
content. Regarding individual major compounds, a significant decrease in hexanal
content and a contemporary increase in benzaldehyde were recorded in the samples
treated with oven drying after sun drying.
Storage is also an important effect on the volatiles of figs. Villalobos et al. (2018)
studied the effect of modified atmosphere packaging (MAP) on the volatiles of two
fig cultivars ‘Cuello Dama Negro’ (CDN) and ‘San Antonio’ (SA) during post-
harvest storage. CDN exhibited slight changes in the volatiles of the control and
22 Fig Volatiles 519

modified atmosphere-packaged samples during cold storage at 0 °C. However, the

volatiles of cultivar SA was affected by cold storage and packaging. An increase
was observed in esters, pyranones, and aldehydes during the cold storage period of
control samples. In addition, the ethyl acetate, 3-methyl butanal, and benzaldehyde
were largely influenced by the cold storage. In their study, the microperforated M50
film for the cultivar SA preserved volatile compounds of figs without negatively
affecting the flavor of the figs.

5 Biological Activities of Fig Volatiles

Biological compounds in foods possess different functions, such as antioxidant and

antimicrobial effects. Because essential oils are rich in such compounds, they play
an important role in protecting human health.
Nafis et al. (2019) evaluated essential oil’s antioxidant and antimicrobial activi-
ties from fig leaves. The results from three antioxidant tests (DPPH· radical scav-
enging, β-carotene acid bleaching test, and reducing power assay) showed that
essential oil had moderate antioxidant activity (IC50 values from DPPH·, reducing
power, and β-carotene/linoleic acid assay were 1.45 mg/mL, 1.92 mg/mL and
1.56 mg/mL, respectively). Like antioxidant activity, the essential oil of figs also
exhibited moderate antimicrobial activity. Yeast strains and Gram-positive bacteria
were generally more sensitive to the essential oil when compared to Gram-negative
bacteria.

6 Conclusion

Fig (Ficus carica L.) is an important fruit grown in many countries with moderate
climates. Such countries as Turkey, Morocco, and Egypt are the major producers of
fig fruit. Since the fruit contains many bioactive components having different activi-
ties such as antimicrobial, antioxidant, anti-inflammatory, anticarcinogenic, and
antidiabetic, the consumption of the fresh fig plays an important role in preserving
human health. In addition to the fresh fruit, the essential oil obtained from figs is
also a rich source of many bioactive components. Several studies revealed that fig
essential oil is a mixture of different chemical components, making the essential oil
a complex extract with numerous health-promoting impacts. The composition of the
fig essential oil depends on the part of the fruit and the extraction method used in
essential fig oil production. The ongoing studies will enlighten different bioactive
effects of figs, and consecutively the worldwide production and the consumption of
this beneficial fruit will increase.
520 M. Kiralan et al.

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Chapter 23
Fig Enzymes: Characterization, Biological
Roles, and Applications

Hesham A. El Enshasy, Bassam Abomoelak, Roshanida A. Rahman,

Ong Mei Leng, Dalia Sukmawati, and Zaitul Iffa Rasid

1 Introduction

Fig (Ficus carica) is an important source of many functional nutrients supporting

human health. Therefore, it was considered a superfood/nutrient-dense and energy-
dense food (Trad et al., 2014; Caliskan, 2015; Khadhraoui et al., 2019). This is
based on its high contents of sugar, fiber, organic acids, micronutrients, and other
biofunctional molecules such as polysaccharides, phenolics, flavonoids, volatile
compounds, and many other molecules which play a significant role in fig taste,
fragrance, and biological activities (Gibernau et al., 1997; Oliveira et al., 2009;
Chawia et al., 2012; Yang et al., 2015; Wang et al., 2017; Arvaniti et al., 2019; Zhao
et al., 2021). Different studies have reported on the bioactivities of fig fruits and
leave with potential applications as antioxidant, hypoglycemic, antidiabetic, anti-
inflammatory, anthelmintic, antibacterial, antifungal, nematocidal, and antiviral
(Solomon et al., 2006; Liu et al., 2011; Hosainzadegan et al., 2012; Debin et al.,
2018; Yousef et al., 2019; Sieniawska et al., 2022). In addition, fruit extract, leave

H. A. El Enshasy (*) · R. A. Rahman · Z. I. Rasid

Institute of Bioproduct Development, Universiti Teknologi Malaysia, Skudai, Malaysia
e-mail: [email protected]; [email protected]; [email protected]
B. Abomoelak
Arnold Palmer Hospital Pediatric Specialty Diagnostic Laboratory, Orlando, FL, USA
e-mail: [email protected]
O. M. Leng
Harita Go Green Sdn Bhd, Ulu Tiram, Malaysia
D. Sukmawati
Biology Department, Faculty of Mathematics and Natural Sciences, Universitas Negeri
Jakarta, Jakarta, Indonesia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 523
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_23
524 H. A. El Enshasy et al.

extracts, and latex exhibited inhibitory effects on the proliferation of cancer cells in
in vitro study using HeLa, stomach, and esophagus cancer cell lines (Hashemi et al.,
2011; Khodarahami et al., 2011; Amessis-Ouchemoukh et al., 2017; Rahmani and
Aldebasi, 2017; Mopuri et al., 2018). Besides the potential medical applications, fig
leave extract has also been used in food industries as a preservative besides the
potential medical applications. For example, recent research reported on the poten-
tial use of fig leaves extract alone or in combination with olive leaves extract as a
natural preservative of pasteurized buffalo milk which can extend the shelf life with-
out altering the milk property (Abdel-Aziz et al., 2020).
In addition to a large number of research deals with the organic and inorganic
contents of the fruits, other studies reported on the presence of a wide range of
enzymes that play a regulatory and functional role during the development and rip-
ening of many fruits. The overall physiochemical and metabolic profiles of fruits are
highly affected by many environmental factors which exhibit a strong influence on
the performance of the enzyme system and thus affecting the fruit ripening process
(Jain et al., 2001; Abu-Goukh et al., 2010; Bae et al., 2014; Gill et al., 2017; López-
Huertas & Palma, 2020). In fig, different enzyme systems have been reported,
including amylases, proteases, invertases, and lipase (Lazreg-Aref et al., 2018). The
nutritional content and bioactive compound profile of fig fruit are governed by the
activities of the enzyme system in each stage of development, which also deter-
mines the carbohydrate, protein, and organic acid profile (Sedaghat & Rahemi,
2018; Cui et al., 2019). Of different enzymes studied, ficin is considered the most
attractive biocatalyst and one of the well-studied enzymes in fig. This is based on its
unique characteristics and high potential applications in industrial, food, and phar-
maceutical industries.

2 Fig Enzymes

Proteolytic enzymes were the first group reported in the latex of fig tree (Ficus
carica L). The early studies were focused on the seasonal variation effect on the
enzyme content of latex and initial kinetic characterization for the optimal pH of the
enzyme (Robbins, 1935a, b). Since then, much research has been focused on the
enzyme content of fig latex in different plant parts as an attractive natural resource
for enzymes of diverse industrial applications.
Figs include different types of enzymes that play a regulatory role during fruit
development, growth, and ripening. However, amylase (both of α- and β-types),
protease, peroxidase, invertase, and lipases are the main reported enzymes (Lazreg-
Aref et al., 2018; Sedaghat and Rahemi, 2018). In general, the chemical and enzy-
matic profile of the fig latex are varied and influenced mainly by the type of cultivar,
harvesting time, and the maturity stage (Lazreg-Aref et al., 2017, 2018). The fig
fruit ripening process is carried out as a function of enzyme-mediated cell wall
hydrolysis, especially polysaccharides. Both α- and β-amylase activities increased
significantly up to the ripening phase and decreased gradually once the fruit entered
23 Fig Enzymes: Characterization, Biological Roles, and Applications 525

the senescence stage. This plays a regulatory role in the starch conversion to sugar
during the ripening stage (Sedaghat and Rahemi, 2018).
On the other hand, it can also be assumed that the invertase enzyme could play a
regulatory role in determining the developed fruit‘s metabolic profile and glucose/
fructose ratio. Of different enzymes present in the fig latex, proteases exhibited the
highest activity compared to amylase and lipase in almost all types of studied fig
cultivars. It has also been reported that, for most of the fig cultivars studied, the
highest latex content and maximal protease activity have been reported in fig fruits
collected from the mid-maturity to the early ripe stage. However, some fig cultivars
exhibited the highest protease activity at the fully ripe stage (Lazreg-Aref et al.,
2017). On the other hand, lipase activity is almost absent at the unripe fruit stage in
most cultivars, and the increase of lipase activity was parallel to the increase of
protease activity during the receptivity period. The increase in lipase activity cata-
lyzed the pathway of volatile compound production during the fruit maturity stage,
which significantly attracted more pollinators to improve the quality of produced fig
fruit (Gaaliche et al., 2011). Based on lipase activity, many volatile compounds
considered fingerprints of fig ripening are produced from lipids and fatty acids.
These include ketonic compounds and specific esters like isoamyl acetate and butyl
acetate (Lazreg-Aref et al., 2018).
It is also interesting to note that recent research has been reported on the presence
of enzyme inhibitors fractions in the fruit latex of Ficus carica L. subsp. carica. The
composition of the isolated fraction was determined by LC-MS/MS and showed
that it includes xanthone, organic acid, chromone derivative, fatty acid, and pheno-
lic acid. Furthermore, this semi-purified fraction exhibited an inhibitory effect on
α-amylase and α-glucosidase enzymes (Paşayeva et al., 2020). Thus, it could be
concluded that these inhibitors could play a regulatory role in the enzymatic activi-
ties during the fruit ripening process.
In addition to the previous enzymes, more recently, three Ficus carica peroxi-
dases isoenzymes were isolated from fruit latex and denoted as (FP1, FP2, and
FP3). These enzymes have been isolated using CM-sepharose, DEAE-sepaharose,
and Sephacryl S-200. The isoenzyme FP1 is monomeric with a molecular weight of
30,000 Dalton. Both FP1 and FP3 exhibited almost the same optimal pH and tem-
perature of 5.5 and 40 °C, respectively. On the other hand, the optimal catalytic
activities for PF2 were observed at pH 7.0 and a temperature of 30 °C.
Furthermore, the biocatalytic activities of FP1 and FP2 were found to be increased
by 11% and 53% in the presence of a high calcium concentration of 10 mM, respec-
tively. At the same time, the presence of calcium ions at this concentration decreases
the activity of FP3 by 67%. On the other hand, applying copper ion at a concentra-
tion of 10 mM inhibits the activity of FP1 completely and increases the catalytic
activities of FP2 and FP3 by 9% and 269%, respectively (Elsayed et al., 2018). Based
on the biocatalytic properties of these enzymes in terms of temperature and pH
range, they could have high potential industrial applications. However, the fig latex
and its associated enzymes play a regulatory role in fruit maturity and ripening and
help defend against different plant pathogens, including microbes and insects.
526 H. A. El Enshasy et al.

3 Ficin: The Most Important Enzyme of Fig Latex

Ficin (EC 3.4.22.3; CAS # 9001-33-6), also known as ficain, Debricin, or higuer-
oxyl delabarre, is a nonspecific sulfhydryl protease that could be extracted from the
latex sap of different parts of the plant such as unripe fruit, leaves, and stem. This
enzyme is characterized by the presence of a single reactive cysteine (Cys) at its
active site. The amino acid sequence in the enzyme active site is similar to papain.
Ficin biocatalytic reaction involves three steps. First, the rapid development of loose
substrate-enzyme complex; Second, acylation of the -SH group of the enzyme
active site by the carbonyl group of the substrate; Third, the decomposition of the
acylated enzyme complex to produce the enzyme and product. However, besides the
well-known proteolytic activities of ficin, a recent study also showed this enzyme’s
intrinsic peroxidase-like activity. The enzyme showed the capacity to act as protease
and peroxidase based on the presence of two active sites in each biocatalytic reac-
tion and thus could catalyze more than one kind of substrate (Yang et al., 2017).
The first report of ficin isolation in the pure crystal structure is back in the late
1930s (Walti, 1938). This enzyme was initially reported as an enzymatic mixture
made of several types of cysteine proteases. It can be found in the fig tree latex in
different isoforms. However, the most commonly studied ficins are endophytopep-
tidase which exist in the latex of Ficus carica, Ficus glabrata, and many other spe-
cies. In early research, ficin from Ficus carica was purified, characterized, and
reported as a homogenous enzyme after isolation using SDS-PAGE electrophoresis
and gel filtration chromatography. It is composed of a single polypeptide chain of
molecular weight of 23,100 ± 300 Dalton when determined using the MALDI-TOF
technique, with optimal pH between 6.5 and 8.5 (Devaraj et al., 2008). Another
study also reported that the ficin obtained from Ficus racemose has a molecular
weight of 55,500 Dalton with optimal pH rage 4.5–6.5 at 60 °C (Arshad et al.,
2016). Another study reported three ficin isoforms (A, B, and C) in Ficus carica cv.
Sabz latex (Zare et al., 2013). All ficin isoforms undergo an autolysis process when
stored at high temperatures. However, ficins B and C isoforms were less stable when
stored at cold temperatures than ficin A. It was also reported on the contribution of
a cysteine residue of the active site in the biocatalytic activity of ficin. This has been
proven by losing enzyme activity by adding a cysteine inhibitor. Based on the pub-
lished data in the RCSB Protein Database bank, ficins are present in 5 isoforms
(Fig. 23.1), namely (Ficin A, BI, BII, CI, CII, and D2). The PDB code for the
enzymes given in the database is: for ficin A (4YYQ), for ficin BI (4YYR), for ficin
BII (4YYS), for ficin CI (4YYU), for ficin CII (4YYV), and ficin D2 (4YYW).
However, other research has also reported the presence of a glycosylated proteinase
from Ficus carica var. Höraishi and denoted as ficin S. This enzyme isoform was
electrophoretically homogeneous with optimal pH of 8.0 and optimal temperature
of 60 °C (Sugiura and Sasaki, 1974). The main feature of this enzyme is the wide
range of pH stability between 2 and 8 when stored for 20 h at 4 °C. The activity of
this enzyme increases in the presence of cysteine and mercaptoethanol. However,
23 Fig Enzymes: Characterization, Biological Roles, and Applications 527

Ficin A (4YYQ) characterized by X-Ray Ficin B (4YYR), crystal form I,

diffraction at a resolution of 1.594 Å characterized by X-Ray diffraction at a
resolution of 1.349 Å

Ficin B (4YYS), crystal form II, Ficin C (4YYU), crystal form I,

characterized by X-Ray diffraction at a characterized by X-Ray diffraction at a
resolution of 1.35 Å resolution of 1.18 Å

Ficin C (4YYV), crystal form II, Ficin D2 (4YYW) characterized by X-Ray

Characterized by X-Ray diffraction at a diffraction at a resolution of 1.446 Å
resolution of 1.902 Å

Fig. 23.1 Crystallographic structure of different ficin isoforms isolated from Ficus carica,
N-terminus blue, C-terminus red. (Obtained from RCSB Protein Database bank)

the main differences between ficin S and other isoforms are only in the isoelectric
point and the sugar residue.
However, another study also reported the presence of new isoforms Ficin C III,
and Ficin D is further divided into isoforms D2-I and D2-II (Haesaerts et al., 2015).
528 H. A. El Enshasy et al.

Another research group also claimed the discovery of a new isoform and gave the
name of Ficin E, a homogeneous protease of molecular weight of 24,294 Dalton.
This ficin isoform is characterized by its maximal activity at pH 6.0 and temperate
of 50 °C, making it attractive with high potential applications in many industries
(Baeyens-Volant et al., 2015). However, based on the increased market demand for
ficin, a new attempt has been made to produce this important endoprotease in micro-
bial biofactory. The ficin production gene has been successfully cloned and
expressed using a lactose induction system in Escherichia coli BL21. The produc-
tion medium was optimized, and the bioactive enzyme was produced in a submerged
cultivation system.

4 Industrial Applications of Ficin

Based on its strong proteolytic properties, availability at a reasonable cost, and

attractive biocatalytic properties, ficin has been used as an attractive enzyme candi-
date in many industries. In general, the use of plant enzymes, especially in the food
and pharmaceutical industries, is highly interested based on their safety and ease of
approval as Generally Regarded as Safe (GRAS) biocatalysts according to Food and
Drug Administration (FDA). In addition, the production of ficin from figs can be
carried out using non-edible plant parts such as leaves and stems, which do not
interfere with the food bioresources. These plant parts are also considered wastes in
fig plantations. Based on all these advantages ficins are widely used as biocatalysts
for bioactive compounds production, food, and pharmaceutical industries, with
many new novel applications in medical and cosmetic fields (Morellon-Sterling
et al., 2020). In general, ficin is widely used in free form. However, different immo-
bilization approaches have been applied to increase enzyme efficiency and stability.
Enzyme immobilization improves the ficin kinetics behavior in terms of activity and
stability and decreases the biocatalysts cost in the conversion process based on its
potential reusability in operation when applied in repeated batch or continuous
modes of operation. Figure 23.2 summarizes the main current applications of ficins
in biotechnology industries.

4.1 Bioactive Compounds Synthesis

Ficin is a cysteine endopeptidase enzyme with a highly efficient capacity to break

protein. It was able to cleave protein at the glycosylic side of Gly, Ser, Thri, Met,
Lys, Arg, Tyr, Ala, Asn, and Val. Thus, it can be widely used in peptide hydrolysis
to produce a low molecular weight-specific polypeptide chain. Therefore, milk pro-
tein hydrolysate of shorter peptide chain production for infant feeding is one of the
23 Fig Enzymes: Characterization, Biological Roles, and Applications 529

Fig. 23.2 Different applications of ficin in biotechnology industries

main fields of the current ficin applications. It is well known that cow milk is unsuit-
able for infant feeding because of the presence of large casein micelles that infants
are not digestible. In addition, the presence of casein and calcium in high concentra-
tions can inhibit the absorption of dietary non-heme iron, affecting iron nutrition
(Ziegler, 2007). Therefore, for the development of infant milk, enzymatic hydroly-
sis is necessary to produce high bioavailable nutrients of high nutritional value and
low allergic effects (Xie et al., 2013). However, milk protein hydrolysis can produce
hydrolysate of various molecular weight peptides with low/zero free amino acid
content or a high amount of free amino acids with short-chain peptides (depending
on the enzyme used). When ficin is applied in this process, it produces a highly
hydrolyzed protein with a significant reduction of allergenicity based on the reduc-
tion of epitopes concentration which IgG and IgE can recognize in the infant
immune system (Verhoeckx et al., 2015). In another research, ficin was able to
hydrolyze bovine casein and produced eight peptides of antioxidant properties (De
Pierro et al., 2014). The main advantage of ficin over pepsin in this process is the
ability of ficin to hydrolyze protein under neutral and alkaline conditions
(Aider, 2021).
Ficin has also been reported as the most suitable protease to produce a high yield
of active bivalent F(ab)2 fragment from the murine monoclonal IgG1 with high
quality suitable for clinical trials. Therefore, it has high potential in the antibody
production industry as a safe and specific protease (Mariant et al., 1991).
530 H. A. El Enshasy et al.

4.2 Food Industries

Plant enzymes have been widely accepted in food processing in many cultures. Of
different plant proteases used, Papain (from Papaya fruit), Bromelain (from
Pineapple fruit), actinidin (from Kiwi), and ficin (from Fig) have a long successful
history in food industries (Troncoso et al., 2022). Ficin, in particular, is often used
to improve the properties and texture of meat, prepare cheese, clarify beverages, and
prevent chill haze. In meat industries, ficin has been used as a tenderizer alone or in
a combination of other plant proteases such as papain or bromelain.
In beverage industries, fig enzyme drink is manufactured using a fermentation
process by adding easily fermented sugars such as glucose or sucrose to speed the
fermentation process. As fig is microbial-rich fruit, the added sugar is fermented
ethanol and carbon dioxide directly without any exogenous addition of any starter
culture. The product of this fermentation process is acidic and composed of a mix-
ture of fig syrup and fig vinegar with a more sourish taste if left for a longer fermen-
tation time. It has been claimed that the fig enzyme-rich drink can provide many
functional roles in the human body by promoting digestion, helping to regulate
blood pressure and sugar level, reducing cholesterol, and reducing the cancer risk
due to the presence of many antioxidants. In particular, ficin is also used to hydro-
lyze protein in beer to decrease the turbidity developed during cold storage.
In meat industries, ficin is considered among the best plant enzymes in the ten-
derization process when used alone or in combination with other enzymes
(Madhusankha & Thilakarathna, 2021). This is based on its ability to hydrolyze the
muscle protein and collagen to soften the meat tissue. Ficin is among the best suit-
able for collagen and elastin degradation at an acceptable range for fresh meat pro-
cessing (pH 5–8) and (temperature 45–45 °C) (Tántamacharik et al., 2018).
Therefore, the solubility of meat proteins can be increased after ficin marination. In
addition, ficin is able to degrade elastin at low temperatures with less reactivity with
myofibrillar proteins and collagen (Singh et al., 2019). It has also been used in the
sausage industry as it has been used for the efficient cleaning of animal intestines.
Another study also reported that adding ficin to soy proteins helps to improve the
final sausage quality and customer acceptability (Ramezani et al., 2003). In seafood
industries, ficin was added alone or in mixtures of other enzymes such as amylase
and cellulase shrimp processing industries to ease the separation of the shell from
the meat (Venugopal et al., 2000; Fernandes, 2016).
In the Cheese industry, ficin help in the milk coagulation process, which is
important for milk clotting in dairy industries. It has been reported that plant prote-
ases such as ficin and papain had a more significant effect on crud milk proteolysis
than calf rennin. Other studies also reported that ficin could improve ewes milk crud
by producing less viscous with a better compact structure (Shabani et al., 2018). It
has also been reported that adding ficin to ovine milk can produce a better-coagulated
product with a higher yield than traditionally used chymosin (Bornaz et al., 2010).
23 Fig Enzymes: Characterization, Biological Roles, and Applications 531

4.3 Pharmaceutical, Medical and Cosmetic Industries

In general, plant proteases have a wide range of applications in pharmaceutical,

medical, and cosmetic industries based on their unique enzyme behaviors in terms
of unique substrate specificity, high stability, the capacity to function at a wide
range of pH, and the ability to work at relatively higher temperature (Balakireva
et al., 2019). Of different plant proteases, ficin can act as an antiparasitic agent
based on its high proteolytic activity with the capacity to digest parasite eggs and
mature worms (Behnke et al., 2008). Ficin has high potential when used alone or in
combination with papaya latex to act as antiparasitic against the adult worms of
Heligmosomoides bakeri. Clear damage was observed to the worm cuticle when
exposed for 30 min to the saline solution of only 30 μm ficin or 200 μm crude
papaya latex or a combination thereof (Stepek et al., 2007). Ficin was also success-
fully used as a hemostatic agent. It can reduce the prothrombin time of normal
plasma or coagulation factor deficient plasma, activate partial thromboplastin, and
act as a factor X activator. This effect is mediated through the successive hydrolysis
in the heavy chain between Leu178 and Asp 179, Arg 187 and Gly 188, Arg 194 and
Ile 195, and the release of a carboxy-terminal peptide (Richter et al., 2002; Morellon-
Sterling et al., 2020). On the other hand, recent research reported the presence of
fibrinolytic serine protease in fig latex which can control the ficin coagulation stim-
ulatory effect (Hamed et al., 2020).
Ficin has also been applied in skin treatment based on its activity as an inhibitor
of skin inflammation. This is mediated through the inhibition of mRNA and protein
expression level of inducible NO synthase and inhibition of the phosphorylation of
inflammatory signal proteins (Cho et al., 2019). The same study also showed the
potential use of ficin in cosmetics as a whitening agent as it can act as a melanin
pigment inhibitor using a mouse melanoma cell model. Another exciting study
reported the high potential application of free and chitosan-immobilized ficin as an
antibacterial agent with wound healing properties. Adding ficin in the free or immo-
bilized form in the rat in vivo model reduced the skin wound area caused by
Staphylococcus aureus twofold after 4 days of treatment instead of 6 days. In addi-
tion, the application of immobilized enzymes reduced the cell count of S. aureus by
3-log on the wound surface after only 6 days compared to 10 days in untreated skin
(Baidamshina et al., 2020). This clearly shows the high potential use of chitosan
immobilized ficin as part of wound-dressing materials to accelerate wound healing
and prevent contamination during the healing process.
Other interesting research also reported the potential use of ficin in cell banking
of camel semen samples (Niasari-Naslaji et al., 2016). In this research, ficin has
decreased camel semen viscosity, limiting the processing and preservation process.
However, adding ficin in a concentration of 0.05 mg/ml followed by stirring for
5 min can decrease the semen viscosity without adversely affecting sperm viability.
In immunohistochemical research, ficin was also applied as a potential trypsin
substitution and ease antigen site exposure during the pre-treatment phase before
the aldehyde fixation process (Taschini et al., 1987). In clinical diagnosis industries,
532 H. A. El Enshasy et al.

ficin has been applied to increase the sensitivity of in vitro diagnostic tests (IVD).
An early study showed that using ficin-treated cells in a serological test of blood
grouping can improve the blood group’s determination by enabling antibody detec-
tion (Giles, 1960). In another study, in testing the plasma using ficin-treated reagents,
the enzyme proteolytic activities increase the sensitivity of the assay, assisting in
better identification of the antibody in ABO, Rh, Kidd, Lewis P, and I antigen in the
blood group system. On the other hand, ficin inhibit Duffy and MNS system anti-
gens. Thus, adding ficin to the diagnosis process allowed to identify 25 new alloan-
tibodies that were masked and could not be detected without enzyme addition (Hill
et al., 2016).
Recent research also reported on the high potential application of ficin as an
efficient protease enzyme for biofilm removal. It is widely known that microbial
biofilm formation creates problems in treating many microbial infections. The bio-
film creates an extra protection layer of a complex structure that prevent the direct
exposure of microbes to antibiotics and thus creates a specific sort of microbial
resistance. Ficin has been effectively used to disrupt skin biofilm matrix made of
Staphylococcus aureus and Staphylococcus epidermidis. The presence of ficin in
the treatment protocol leads to a twofold decrease in the concentration of the anti-
infectives (ciprofloxacin and benzalkonium chloride) used in skin treatment without
any cytotoxic activity (Baidamshina et al., 2017). Recent research also reports the
high potential biofilm suppressant application when used in concentrations as low
as 4 mg/mL. At this concentration, it exhibited a significant inhibitory effect on
Streptococcus mutans biofilm formation and virulence mechanism involving lactic
acid production and exopolysaccharide synthesis. Therefore, ficin could be used as
an anticaries agent in dental treatment (Sun et al., 2021).

5 Ficin Immobilization to Improve Industrial Applications

Most research on ficin applications in the industry has been carried out using a free
enzyme system. However, in recent years, many studies focused on the immobiliza-
tion of this enzyme to increase stability and reusability and decrease the enzyme
cost in many industrial processes. One of the early attempts was carried out for ficin
immobilization using diatomaceous earth (Celite) as an adsorbent. The immobilized
enzyme was used in traditional Turkish cheese (Teleme) production. As a result, the
optimal temperature was increased from 60 °C, both free and immobilized enzymes
exhibited high milk clotting activity, and the produced cheese by immobilized
enzyme gave better chemical and sensory properties compared to a free enzyme
(Fadýloğlu, 2001). In another research effort, ficin was covalently immobilized on
poly (γ-methyl-D-glutamate) fibers and produced a highly active and stable enzyme
suitable which is more suitable for continuous operation (Jokei et al., 1999). Another
study also reported the successful immobilization of ficin using polyvinyl alcohol
(PVA), with high retention of enzyme activity up to 92%, and was used successfully
for 9 cycles without losses in biocatalytic activities (Rojas-Mercado et al., 2018). In
23 Fig Enzymes: Characterization, Biological Roles, and Applications 533

other research, glyoxyl-agarose was successfully used as an immobilization matrix.

The immobilized enzyme exhibited higher stability, a wider range of pH and tem-
perature activity curves, and catalyzed casein hydrolysis five times without losing
its initial activity (Siar et al., 2017). Another recent study also reported the success-
ful use of immobilized ficin using the same matrix to improve the milk coagulation
process by reducing the clotting time, producing a higher coagulation yield than the
free enzyme (Siar et al., 2020).

6 Conclusion and Future Perspectives

In addition, with the increased awareness of the importance of natural enzymes in

human health, plant enzymes have become more widely used in the food and phar-
maceutical industries. So far, ficin has exhibited unique characteristics as a dual
active enzyme of both protease and peroxidase activities. Therefore, we expect an
increase in its application range, especially in the medical field as a high-end market
for plant enzymes. This is because edible plant parts produce it, considered one of
the safest natural resources according to the FDA. Every year, ficin applications
increase in many industries. It is expected that improving enzyme kinetics using
protein engineering techniques and immobilization methods will increase the
acceptability and allow the penetration of ficin to many industries, replacing in large
part the currently used chemical catalysts. To cope with the increased demand, new
suspension culture cultivation technology in large-scale bioreactors produces a
high-quality enzyme in a shorter time. In addition, this process is not affected by
environmental variables or climate changes. In addition, with the increased knowl-
edge about molecular biology and biosynthesis of ficin enzymes, the ficin produc-
tion genes were recently expressed in microbial biofactory. Therefore, we expect to
see a large-scale production process of ficin using the microbial system shortly.
Further research in all these directions will reduce the production cost of ficin and
make it available to replace other endoproteases in the market.

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Chapter 24
Preventive Roles of Phytochemicals
from Ficus carica in Diabetes and Its
Secondary Complications

Additiya Paramanya, Nimisha Patel, Dinesh Kumar, Fatima Zahra Kamal,

Belkıs Muca Yiğit, Priya Sundarrajan, Prairna Balyan, Johra Khan,
and Ahmad Ali

1 Introduction

Ficus carica L. (Moraceae), or the common fig, is the most important member of
the 23 species of the genus Ficus. It is one of the first plants cultivated by humans
and is native to the Mediterranean and a few parts of Asia (Mawa et al., 2013).
Previously, only the fruit of the plant, i.e., the fig, was consumed only for nutritional
purposes due to its high vitamin and mineral content, while the bark, latex, and roots
were used for topical applications in traditional medicine (Misbah et al., 2013).

A. Paramanya · D. Kumar · P. Balyan · A. Ali (*)

Department of Life Sciences, University of Mumbai, Mumbai, India
e-mail: [email protected]
N. Patel
Department of Life Sciences, J C Bose University of Science and Technology, YMCA,
Faridabad, Haryana, India
F. Z. Kamal
Faculty of Sciences and Techniques, Laboratory of Physical Chemistry of Processes
and Materials, Hassan First University, Settat, Morocco
B. M. Yiğit
Vocational School of Technical Sciences, Department of Forestry, Igdir University,
Igdir, Turkey
P. Sundarrajan
Department of Life Science and Biochemistry, St. Xavier’s College (Autonomous),
Mumbai, India
J. Khan
Department of Medical Laboratory Sciences, College of Applied Medical Sciences,
Majmaah University, Al Majmaah, Saudi Arabia

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 539
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_24
540 A. Paramanya et al.

Figs are an excellent source of proanthocyanidins, a powerful antioxidant with

potential antidiabetic effects. The fruit’s anthocyanin content depends on its skin
color, ranging from yellow, green, violet, purple, and black. These pigments are
found to be highest in black-colored figs and are proven efficient in reducing oxida-
tive stress, inflammation, and carcinogenic effects (Tall et al., 2004; Ercisli
et al., 2012).
Oxidative stress is one of the major contributors to diabetes, antioxidants in fig
might prove beneficial in combating it. Organic extracts of fruits and leaves have
demonstrated induction of insulin secretion and inhibition of enzyme levels such as
pancreatic lipase, α-glucosidase, and α-amylase (Irudayaraj et al., 2016; Mopuri
et al., 2018). In addition, the plants have been proven to show varying degrees of
hypoglycemic activity, suggesting that their use can greatly reduce the effects and
complications of diabetes.

2 Management of Diabetes

Diabetes, commonly known as diabetes mellitus, is a chronic metabolic condition

marked by a shortage of insulin, a high blood glucose level, and insulin resistance.
Diabetes develops when the body’s insulin production is very low or non-existent or
when the body’s response to the released insulin is inadequate (Shoback & Gardner,
2011). Type 1, type 2, and gestational diabetes are the three most common kinds of
diabetes. Type 1 diabetes, also known as insulin-dependent diabetes mellitus
(IDDM) or juvenile diabetes, occurs when the pancreas fails to produce enough
insulin due to beta cell loss (Norman & Henry, 2015). Type 2 is a condition wherein
the body fails to give insulin response appropriately; it is commonly known as non-
insulin-dependent diabetes mellitus (NIDDM) or adult-onset diabetes. Type 2 dia-
betes, also known as non-insulin-dependent diabetes mellitus (NIDDM) or
adult-onset diabetes, is a condition in which the body fails to respond to insulin
adequately. Lack of exercise and weight increase is typical causes (Moller, 2001).
The third type of diabetes is gestational diabetes, which occurs when a pregnant
woman who has never had diabetes gets excessive blood sugar levels, which usually
resolve itself once the baby is born (Coustan, 1993).
Diabetes mellitus has such modest symptoms that around eight million people
are unaware they have it. Frequent urination, hazy vision, irritation, increased hun-
ger and thirst, and weight loss are common symptoms (Cooke & Plotnick, 2008).
The glycated hemoglobin (A1C) test, which indicates the sugar content in the blood
bound to hemoglobin, is one of the diagnostic tests for type 1 diabetes. The goal of
diabetes management is to maintain a normal blood sugar level (Fig. 24.1). This can
be accomplished using appropriate drugs, exercise, dietary changes, and other
methods (Nathan et al., 2005). A sedentary lifestyle is one of the most prominent
risk factors for type 2 diabetes. To keep blood sugar levels under control, it is critical
to improving your lifestyle by exercising and making dietary changes. In a research
study, patients who exercised moderately for 50 min three times a week saw their
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 541

Weight
Control

Timely Healthy
diagnosis lifestyle

Diabetes
Treatment
Regular
Exercise medica-
tions

Gene Insulin
therapy shots

Fig. 24.1 Strategies to treat diabetes

HbA1c levels drop by 0.75% after 8 weeks (Boulé et al., 2001). Diabetes raises the
risk of cardiovascular disease and stroke. Weight loss combined with a healthy diet,
such as a low-carbohydrate diet or a diet rich in fruits, vegetables, whole grains,
low-fat dairy products, and artificial sweeteners, has been shown to help manage
hyperglycemia (Evert et al., 2019).
Diabetes can be treated with a variety of medications. Insulin shots, preferably
long-acting insulin to match the basal rate, are given subcutaneously in the case of
type 1 diabetes. In the case of type 2 diabetes, medications such as metformin are
commonly taken orally Metformin is the first-line treatment for T2DM and gesta-
tional diabetes that works by lowering glucose production in the liver. It reduces
calorie intake by lowering glucose synthesis in the liver, increasing insulin sensitiv-
ity, and exerting anorexiant effects (Krentz & Bailey, 2005). Other anti-diabetic
medications include thiazolidinediones (TZDs), alpha-glucosidase inhibitors,
SGLT2 inhibitors, etc. TZDs lower blood glucose levels by improving insulin sen-
sitivity in cells. Glucosidase inhibitors work by blocking the enzyme alpha-
glucosidase, which delays carbohydrate absorption in the intestines. SGLT2
inhibitors prevent glucose from reabsorbing in the kidneys, resulting in increased
secretion and lower blood sugar levels (Wang & Perri, 2018). In recent years, nano-
technology has become increasingly essential in treating diabetes, with glucose sen-
sors delivering insulin to the desired location. A microcapsule containing auxiliary
542 A. Paramanya et al.

islets of Langerhans cells is implanted beneath the skin of patients in this procedure,
which aids in the restoration of the body’s glucose regulator feedback loop without
the use of severe immunosuppressants (DiSanto et al., 2015).
Gene therapies that use non-viral and viral transduction could help people with
diabetes by reducing islet cell destruction or replacing the insulin gene. Gene ther-
apy could be done in epithelial cells, hepatocytes, or stem cells in vivo or in vitro.
Islet allotransplantation could also restore normal blood glucose levels and prevent
graft rejection (Wong et al., 2010).

3 Glycation

The word glycation has become a well-known term nowadays in biological research.
The glycation process has involved the non-enzymatic interaction of amine groups
of nitrogen-containing compounds viz. proteins, DNA, etc., with the carbonyl group
of reactive sugars. Maillard first described the reaction of glycation interaction in
1912. This reaction is multistep and requires days, weeks, or months to complete
(Yeh et al., 2017).
Heating and storage have enhanced the Maillard reaction during the processing
of food. It leads to unstable Schiff’s base and Amadori products, mainly fructos-
amine, in the early stage of the glycation process. In the intermediate stage of glyca-
tion, these unstable products get converted either into stable adducts or reversed.
The redox reactions, precipitation, aggregation and various other physiological pro-
cesses play a major role in the further generation of advanced glycation end-products
(AGEs), which are considered late-stage glycation products (Kikuchi et al., 2003;
Kumar et al., 2021). AGEs are known as irreversible and deleterious products. The
reactivity of sugars plays an important role in increasing glycation. The monocar-
bonyl sugars, such as glucose, fructose, etc., were less reactive than the dicarbonyl
species, viz. methylglyoxal. The generated AGEs in glycation observed their inter-
ference in diabetes secondary complications, neurodegenerative disorders, and car-
diovascular disorders (Jha et al., 2018).
The generated AGEs are classified into fluorescent crosslinking AGEs, nonfluo-
rescent crosslinking AGEs, and nonfluorescent non-crosslinking AGEs. The
crossline, pentosidine imidazolium dilysine crosslinks, N-ɛ-carboxyethyllysine
(CEL), and N-ɛ-carboxymethyl-lysine (CML) are the few well-characterized AGEs
generated in the process of glycation. Most AGEs are irreversible in their produc-
tion except for pyrraline and pentosidine (Yeh et al., 2017). In the last few decades,
scientists have been dedicated to developing therapeutic drugs/compounds that
could hinder or stop the production of AGEs. For example, aminoguanidine, a syn-
thetic drug, is widely researched for inhibiting the glycation process. In addition,
the natural compounds and secondary metabolites are also explored for the treat-
ment of glycation-mediated complications and inhibition of the generation of
glycation-mediated products in either stage of glycation (Abbas et al., 2015).
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 543

4 Oxidative Stress

Oxidative stress corresponds to the systemic inefficiency in balancing the level of

reactive oxygen species produced in cells due to normal metabolic activities (Zahra
et al., 2021). This imbalance affects cells’ redox state, compromising their defense
mechanisms to detoxify the reactive species (Nita & Grzybowski, 2016). In addi-
tion, prolonged oxidative stress causes damage to all components of the cell, includ-
ing proteins, lipids, and DNA (Kamal et al., 2022).
Because of their high content of unsaturated fatty acids, lipid structures, particu-
larly cell membranes, are vulnerable to (ROS/RNS) assault (Zahra et al., 2021). The
oxidative lipid damage leads to what is known as “lipid peroxidation”. It has been
found that lipid peroxidation causes oxidative changes in the structure, function,
and permeability of the biological membranes (Niki, 2008; Zahra et al., 2021). It
also produces potentially toxic byproducts like malondialdehyde (MDA),
4-hydroxy- 2-nonenal (HNE), and reactive carbonyl species (RCS), which are
chemically reactive and covalently modify vital macromolecules such as DNA
bases, proteins and low-density lipoproteins (LDL) leading to cytotoxic events
(Niki, 2008; Zahra et al., 2021).
DNA is also influenced by oxidative stress. Oxidative stress causes additional
DNA damage, including mismatch, inter-strand crosslink, single-strand breaks,
intra-strand crosslink, oxidation of specific bases, and double-strand breaks
(Fig. 24.2). Inefficient DNA repair mechanisms cause genome instability, which

Fig. 24.2 DNA oxidative damage

544 A. Paramanya et al.

leads to altered cell behavior or cell death (Alhmoud et al., 2020). The base altera-
tion of 8-oxoguanine is one of the most important byproducts of DNA oxidation
(8-oxo-G) (Zahra et al., 2021). It is mildly mutagenic and causes a G: C to T: A
translocation (Zahra et al., 2021). It has been linked to cell transformation and can-
cer initiation (Zahra et al., 2021). Proteins have not escaped oxidative damage. In
fact, because of their high concentrations, they are directly targeted by ROS/RNS
(Niki, 2008). Deamination, decarboxylation, conformation modifications, changes
in electrical charges, cross-linking, and side-chain modification may all occur as a
result of protein oxidation, resulting in to halt of enzyme activity and accumulation
within cells and the extracellular environment (Niki, 2008; Zahra et al., 2021). For
example, histidine can be modified to oxo-histidine, “a damaged form of histidine”
after exposure to oxidative stress (Ezraty et al., 2017). As a result of its toxic insult
to biomolecules, oxidative stress can lead to grave diseases.
The accumulation of oxidative stress is the primary cause of aging on a cellular
level (Romano et al., 2010). It is also associated with metabolic diseases like diabe-
tes, neurodegenerative diseases like Parkinson’s, Alzheimer’s, depression, and car-
diac diseases like myocardial infarction and atherosclerosis (Roberts & Sindhu,
2009). Besides, several diseases like cancer, vitiligo, and increased severity of
microbial infections have been traced to oxidative stress (Chakraborti et al., 2019).
Although the generation of free radicals, as a metabolic process, is the primary
cause of oxidative stress, they do not always induce the same. In their production,
they are instantly neutralized with the help of antioxidants (Kamal et al., 2022).
However, stress, chemical exposure, and genetic/environmental factors more exten-
sively trigger oxidative stress mechanisms. Therefore, the effects do not show path-
ological symptoms instantly (Zahra et al., 2021). Instead, they accumulate silently,
causing extensive damage to macromolecules resulting in dreadful outcomes like
cancer, aging, and death (Burton & Jauniaux, 2011; Preiser, 2012).

5 Ficus Species and Its Phytochemicals

Ficus carica L. is an economical and valuable plant. Fig has many usage areas.
Fruits of figs are valuable as food and are used in the medicinal, getting paint, cos-
metic, and medicine industries. Apart from these, figs are also used to produce raki
and animal feed. (Kavak & Önal, 2006; Kıyak, 2009; Özbek, 1978). Figs is a plant
considered sacred in religion and dates back to ancient times. In addition, the
Hittites used these plants and recorded their names in their texts. Fig is among the
trees recorded in Hittite cuneiform texts. It is known that figs were used during the
making of thick sacrificial bread in these texts (Ünar, 2019). F. carica has a lot of
chemical components. For example, fruits and leaves have Anthocyanin, flavonol,
flavones, phenolic acid, volatile, aldehydes, alcohols, ketones, esters, monoterpens,
sesquiterpens, norisoprenoids, miscellaneous compounds (Fig. 24.3).
At that time, leaves and roots have coumarin and sterol. F. carica latex is special
latex. In addition, it has aldehydes (pentanal, hexanal, heptanal, and benzaldehyde),
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 545

Whole fruit
Catechin
Fruit peel
Rutin
Epicatechin
Fruit pulp
Galic acid
Cyanidin-3-O-
Chlorogenic acid
Leaf Extract
Quercetin-o- rutinoside
gluconicide Pelargonidin-3-O-
Luteolin-7- O- (+)-catechin
Quercetin-3-O- rutinoside
Glucoside Caffeoylmalic acid
malonyl-galactoside Carboxypyrano-Cy 3-
Quercetin Caf feic acid
rutinoside
Ellagic acid Kaempferol derivative
Cyanidin-33-
malonylglycosyl-5- Quercetin derivative
glucoside
Bergapten

Fig. 24.3 Phytochemicals from different parts of F. carica plant

alcohols (1-butanol-3-methyl, 1-butanol-2-methyl, 1-pentanol, 1-hexanol,

1-heptanol, phenylethyl alcohol, phenylpropyl alcohol), Ketones (6-methyl-5-
hepten-2-one), monoterpenes (α-thujene, α-pinene, β-pinene, limonene, eucalyptol,
terpinolene, cis-linalool oxide, linalool, epoxylinalol), sesquiterpenes (α-guaiene,
α-bourbonene, β-caryophyllene, trans-α-bergamotene, α-caryophyllene,
s-muurolene, germacrene D, cadinene, a-calacorene), miscellaneous (methyl salicy-
late, quinoline, psoralene), organic acids (oxalic, citric, malic, quinic, shikimic,
fumaric), fatty acids (myristic, pentadecanoic, palmitic, heptadecanoic, cis-10-
heptadecenoic, stearic, oleic, elaidic, linoleic, arachidic, heneicosanoic, behenic,
tricosanoic, lignoceric), steroids and amino acids (Barolo et al., 2014; Duman &
Yazıcı, 2018). When the studies have been examined, it is common to find studies
on the fruit and latex properties of figs. However, it is possible to come across stud-
ies on the wood of the fig plant in recent years. Moreover, the use of fig plants for
landscaping has been emphasized in the research (Serin & Penezoğlu, 2020).

6 Natural Products from Ficus Species

as Antidiabetic Agents

Various species of Ficus provide various valuable products such as edible fruits,
fodder, and fuel wood, traditionally used as religious, ornamental, and medicinal
plants by humans. It has been globally documented that the Ficus plants are signifi-
cantly spiritual and provide material resources for humans (Shi et al., 2014). In the
genus of Ficus, there are as many as 18 species, of which the species having impor-
tance in traditional medicines are Ficus benghalensis, popularly known as the Indian
546 A. Paramanya et al.

Banyan tree, Ficus hispida is known as opposite leaf fig, Ficus elastica, also called
rubber fig, Ficus religiosa which is also popularly known as peepal tree and Ficus
carica also referred to as Figs or locally as Anjir in India. Among the different plant
species, one of the prominent and frequently used plants in folk medicines like
Siddha, Unani, homoeopathy, and Ayurveda, as well as used for the treatment of
diabetes, is Ficus benghalensis (Kumari et al., 2016) as well as Ficus carica.
Pharmacological studies support the ethnomedicinal uses of various Ficus species,
which have been found to have anti-inflammatory, anticancer, and antidiabetic prop-
erties (Lansky et al., 2008). A wide range of pharmacological properties, both in
vitro and in vivo, have been exhibited by F. carica. Some of these properties are
anticonvulsant, anti-inflammatory, antiviral, antitumor, antidiabetic, antimicrobial,
analgesic, antioxidant, antiulcer, antiasthmatic, antianxiety, antihypertensive,
anthelmintic, apoptosis inducer, estrogenic, endothelin receptor antagonist, hypo-
lipidemic, cognitive enhancing, and immunomodulatory properties (Kurniawan &
Yusuf, 2021).
The indigenous people worldwide have traditionally been using the different
parts of various Ficus species to cure various diseases, including diabetes. Several
studies using in vivo and in vitro models have reported that the extracts are made
from different parts of Ficus species (particularly F. carica, F. benghalensis, F. del-
toidei, F. glumosa, F. religiosa, and F. racemosa), have potent anti-diabetic proper-
ties. Traditionally the fruits, seeds, leaves, bark, tender shoots, and latex of F. carica
leaves have been used as an anti-inflammatory agent to treat diarrhea, jaundice, and
nutritional anemia (Canal et al., 2000).
In African medicine (Olaokun et al., 2014), Ayurveda (Rangika et al., 2015), and
Siddha medicine, diabetes is treated using the edible fruits and leaves of Ficus car-
ica (Irudayaraj et al., 2017). The antidiabetic effects of organic extracts of different
parts like fruits and leaves of Ficus carica were demonstrated in alloxan- and
streptozotocin-induced diabetic animal models. Inducing insulin secretion, inhibit-
ing absorption and increasing glucose uptake, and inhibiting levels of α-glucosidase,
α-amylase, and pancreatic lipase (Irudayaraj et al., 2016; Mopuri et al., 2018), the
extracts helped lessen any complications arising from diabetes or related conditions.
Antifungal, antioxidant, anthelmintic, antimutagenic, antipyretic, antispasmodic,
hepatoprotective, and hypoglycemic properties have been observed (Vikas &
Vijay, 2011).
To elucidate the treatment of diabetes, the decoction prepared from the bark of
Ficus carica was used (Perez et al., 1996). The bioactive agents believed to be pres-
ent in Ficus are sterols, flavonoids, saponins, polyphenolic compounds, and tannins,
to name a few. Hypoglycaemic activity in rat models was found to be elicited by
ethanolic extracts of leaves of Ficus carica. In addition, the various important bioac-
tive compounds imparting antidiabetic effect in Ficus carica flavonoid compounds,
β-sitosterol, 6-O-acyl-b-D-glucosyl β-sitosterol, and polyphenols are reported to
have antidiabetic effects (Mopuri et al., 2018; Kurniawan & Yusuf, 2021).
Antihyperglycemic activity has been observed in the extracts made from leaves
of Ficus carica (Kurniawan & Yusuf, 2021). In animal models, when ethanol and
ethyl acetate extract of Ficus carica was administered orally, it reduced blood
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 547

glucose levels and increased insulin levels significantly. Fig extract was observed to
correct any decrease in skeletal muscle (a site where glucose uptake is stimulated by
insulin) and hepatic glycogen in diabetes (Irudayaraj et al., 2017). It has been
reported that F. carica fruit extracts lower oxidative stress since oxidative stress is a
major cause of the complications and pathogenesis of type 2 diabetes. The extracts
also modulate the enzymes of antioxidant defense systems, especially α-glucosidase
and α-amylase. Hence, from these studies, the extensive Ayurvedic use of plants
such as Ficus carica for the treatment of diabetes has been corroborated by scien-
tific evidence in recent years.

7 Prevention of Advanced Glycation End Products (AGEs)

by Phytochemicals from Ficus carica

AGEs, a heterogeneous, chemically diverse, and complicated set of chemicals gen-

erated either exogenously or endogenously, have piqued the scientific community’s
interest in recent decades due to mounting evidence of their involvement in various
disorders like diabetes mellitus (Davis et al., 2016). Inhibition of the generation of
ROS compounds during the glycation process; inhibition of the Schiff base pathway
and the production of Amadori products, inhibiting the formation of dicarbonyl
compounds; detoxifying AGE precursors, such as methylglyoxal and influencing
AGEs to bind to their receptors (RAGE) are the fundamental processes that prevent
the formation of AGEs (Gkogkolou & Böhm, 2012).
Clinical trials have been conducted on aminoguanidine, the first chemical to sup-
press AGE production, but it is unlikely to be used for medicinal purposes due to
safety concerns and a lack of efficacy (Dömötör et al., 2022). Therefore, AGE inhib-
itors derived from natural compounds with a low toxicity profile are more attractive
options for the development of functional additives and even medications to cure
and manage diabetes complications and other AGE-related disorders. In addition,
several natural substances with antioxidant characteristics currently show good
inhibitory action against AGE development with minimum toxicity (Song
et al., 2021).
Phytochemical investigations of the plant’s leaves and fruits revealed that they
are high in phenolics, flavonoids, polysaccharides, vitamin C, alkaloids, saponins,
coumarins, tannins, and organic acids, and volatile chemicals, making them a potent
antioxidant (Caliskan & Polat, 2011). Another study demonstrated that the 96%
ethanol extract of F. carica leaf from Indonesia had an IC50 value of less than
1000 μg/ml in the DPPH· assay, indicating it to be effective in suppressing the
development of AGEs compounds (Abdillah And & Prakoso, 2018).
Polyphenol compounds suppress the development of AGEs and provide resis-
tance to the pathogenic effects of AGEs, according to in vitro glycation studies.
Several investigations have found a strong link between antiglycation action and
phenolic extracts (Riyaphan et al., 2021). F. carica is an excellent source of
548 A. Paramanya et al.

phenolic compounds, quercetin rutinoside was the major individual phenolic, and
luteolin (187 mg/kg extract) was the most abundant non-glycosylated flavonoid in
F. carica (Vaya & Mahmood, 2006). An isolated rat heart examined the antioxidant
activity and effects of F. carica leaf extract on ischemia/reperfusion injury. The
treatment groups received enriched solutions containing the extract (0.04, 0.2, and
1 mg/ml, respectively) during stabilization and reperfusion (after 30 min of global
ischemia). The dry sample had a total phenolic content of 12.2 mg GAE/100 g and
flavonoid content of 40.7 mg/g (Allahyari et al., 2014). According to Mujić et al.
(2010), in F. carica, phenolics ranged from 7.24 to 11.1 mg GAE/g dry extract, and
the highest TPC was found in methanolic extracts. The most abundant phenolic
acids and flavonoids in fig; include proanthocyanidins, gallic acid, chlorogenic acid,
ferulic acid, rutin, quercetin-3-O-rutinoside, and epicatechin.
Most of the polysaccharides present in plants have low toxicity; therefore, they
are a valuable source for developing glycation inhibitors, but their mechanism of
action is still unknown. Therefore, the antioxidative activities of water extract and
crude hot-water soluble polysaccharide (PS) from the fruit were investigated, and it
was discovered that both water extract and crude hot-water soluble PS had signifi-
cant scavenging activities at effective concentrations of 0.72 and 0.61 mg/mL
respectively (Yang et al., 2009). Furthermore, it was discovered by Zou and his col-
leagues (2020) that the F. carica PS contains antioxidant and immunity-enhancing
properties. PS, one of the most influential and significant components in F. carica
L., has long been thought to be a favorable immunostimulant that might be exploited
in animal and human immunotherapy (Wang et al., 2022).
F. carica leaves are rich in vitamin C. Vitamin C is a water-soluble vitamin that
preserves the biomolecules while reducing oxidative stress. It makes ionic connec-
tions with proteins, allowing them to bind to them competitively and protect their
physiological structure. In vitro, ascorbic acid can compete with glucose for protein
binding, lowering AGE formation. In diabetic individuals, vitamin C inhibits the
sorbitol pathway and reduces the production of fructose, a highly active sugar in
glycation (Song et al., 2021).
Terpenoids are natural compounds with a wide range of structural and functional
properties. Some terpenoids have been shown to reduce AGE production, for exam-
ple, pentacyclic triterpenoids. They have various biological effects, including anti-
oxidant, anti-atherosclerotic, antihyperglycemic, and anti-inflammatory effects
(Cox-Georgian et al., 2019). Bauerenol, methyl maslinate, lupeol acetate, and olea-
nolic acid are the four triterpenoids that have been identified from the leaves of
F. carica (Mawa et al., 2013). Some terpenoids have excellent antiglycation activi-
ties, but only a few investigations of their antiglycation effects have been conducted,
and no clinical applications have been documented to date.
NMR data confirmed that cyanidin-3-O-rutinoside (C3R) was the main anthocy-
anin in all fruits of F. carica (Solomon et al., 2006). C3R is a naturally occurring
anthocyanin with antioxidant, anti-hyperglycaemic, anti-glycation, and cardiopro-
tective activities. In addition, C3R was found to be a mixed-type competitive inhibi-
tor of pancreatic lipase with an IC50 of 59.4 1.41 M, according to Thilavech and
Adisakwattana (2019).
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 549

Coumarins are heterocyclic chemicals with antiglycative activity that has the
potential to cure insulin-dependent diabetic mellitus. Coumarin was extracted from
the methanol extract of the leaves of F. carica L. using bioassay-guided isolation
(Mawa et al., 2013). Coumarin improves bone turnover in diabetic osteoblasts and
osteoclasts by reducing the production of advanced glycation end products (Pan
et al., 2022).

8 Antioxidants Derived from Ficus in Diabetes Management

The substances that can break free radicals and decrease the oxidative stress found
in living organisms are Antioxidants (Ayinde et al., 2007). These antioxidants can
donate electrons which can inhibit free radical-related oxidation reactions.
Antioxidants are either synthesized endogenously or can be taken from different
naturally occurring sources from plants (Rout et al., 2009; Egbuna et al., 2021).
Different studies are carried out to study the antioxidant property of different parts
of the Ficus species. Researchers reported antioxidant activity against Trolox, fer-
ric, lipid, and lipid peroxidation, reducing power in Ficus plant parts. Some research-
ers also found the free radical scavenging capacity of these antioxidants of the Ficus
plant against DPPH· and 2,2-aninobis (3- ethylbenzothiazoline-6-sulfonic acid) in
dose control studies as the Ficus plant extract contains a high amount of phenolic
components (Mazumder et al., 2009). Ficus plant extract was found to possess anti-
cancer, antidiabetic, Hepatoprotective, antitumor, analgesic, antimicrobial, hypo-
glycemic, gastro protective, anti-inflammatory, antifungal, antimalarial, and
antiparasitic activity (Table 24.1). Most species of Ficus showed antimicrobial
activity (Singh & Goel, 2009).
Different species of Ficus are used as traditional or alternate medicine for dis-
eases including stomachic, hypotension, anti-dysentery drugs, and diabetes. Some
species of Ficus like F.glomerata, F.glumosa, F. carica, F. racemosa, and F. bengha-
lensis are effective in treating diabetes (Li et al., 2021). The aqueous extract of
F. benghalensis is found to regulate insulin secretion and enzymatic activities,
increase insulin secretion, carbohydrate absorption, hepatic glycogen synthesis, and
antioxidant activity in the diabetic patient body (Ren et al., 2022). In vivo study on
male rate found that aqueous extract of F. glumosa, F. carica, F. racemosa, and
F. benghalensis reduces oxidative stress and increases the weight of these rats.
Another study with methanol extract of F. carica leaves helps to control the increase
in lipid peroxide by functioning as a hepatoprotective agent (Kerian, 2021; Sahardi
et al., 2021).
In vitro ethanol extract of fruits of Ficus racemosa Linn. on blood glucose in
experimental animals promote glucose uptake and metabolism or inhibit hepatic
gluconeogenesis since alloxan treatment causes permanent destruction of β-cells.
Furthermore, the DPPH· radical scavenging activity of this extract prevents the ill
effects of diabetes and oxidative stress in vivo studies. Another study by Siddique
and Saleem (2011) on ethanol extract of F. racemosa showed the presence of lupeol
Table 24.1 Antioxidant and biological activities of different parts of plants
550

Species of
Ficus Part of plant Solvent type Antioxidant activity Biological activity Reference
Ficus carica Leaves Hexane, water TEAC: 14.0%, 23.5 acetate/g DW Hepatoprotective activity Mondal et al. (2022)
FRAC: 7.9–16.1 mmol/kg FW with a decrease in lipid
DPPH-RSC: 11.4 mmol/100 g DW peroxides with cytochrome
ABT-RSC: 6.48 mmol/100 g DW p450 complex inhibition
Fruit Dichloromethane ILP: 0.02 mg/mL
N hexane ILP: 1.64 mg/mL
Ficus Stem Methanol DPPH-RSC: 16.2%, ABT-RSC: Hepatoprotective activity by Alzahrani et al. (2021) and
racemosa 8615.3 mmol/g DM reducing the activities of Tewari et al. (2021)
ALT, AST, and ALP
Bark Ethanol DPPH-RSC: 79% Anticancer activity by
reduction of lipid
peroxidation, γ-glutamyl
transpeptidase, and xanthine
oxidase and by generation
of hydrogen peroxide
Hypoglycemic activity by
decreasing blood glucose
level
Roots Water FRAC: 0.5–0.26 mg/mL
Ficus Aerial roots Methanol DPPH-RSC: 71%, ABT-RSC: Antioxidant and Joshi et al. (2021)
benghalensis 10884.6 mmol/g DM hypolipidemic activity by
reduction in lipid
peroxidation, cholesterol
level and triacylglycerol
Acetone, Water FRAC: 0.5–0.26 mg/mL Anticancer and antibacterial Kapile et al. (2022)
activity but no antifungal
activity
A. Paramanya et al.
24

Ficus sur Bark Water TEAC: 14.0%, 23.5 acetate/g DW, Antifungal and antibacterial Sieniawska et al. (2022)
FRAC:7.9–16.1 mmol/kg FW, activities
6.48 mmol/100 g DW, DPPH-RSC:
11.4 mmol/100 g DW, ABT-RSC:
10884 mmol/g DM
Unripe fruit TEAC: 62.3 GAE/g DW,
FRAC:19.6 μmol FSE/mg DW,
DPPH-RSC:7.3 QE/mg DE
Ficus Fruit Water DPPH-RSC: 16.2%, FRAC:0.5– Antitumor activity due to Ghadigaonkar et al. (2021)
religiosa 0.26 mg/mL, ABT-RSC:8615 mmol/g blockage of calcium uptake
DM in pituitary cells
Whole Water, Methanol Antioxidant and antidiabetic Randima et al. (2022)
activity with lowering the
superoxide dismutase
exaggerated activity
Bark Water DPPH-RSC: 79%, ABT- Antimicrobial activity with Murugesu et al. (2021)
RSC:10884.6 μmol/g DM inhibition zone against B.
subtilis
Ficus polita Whole Water TEAC:2.45%, 5.40 mg GAE/g DW Antiviral activity due to Sieniawska et al. (2022)
Vahl inhibition of reverse
transcriptase activity of
HIV-1
Leaves Water Antimalarial action against Ben Nasr et al. (2021)
Preventive Roles of Phytochemicals from Ficus carica in Diabetes…

Plasmodium falciparum
551
552 A. Paramanya et al.

as a super antioxidant (Siddique & Saleem, 2011). Lupeol decreases peroxidation

and is effective against diabetes, cardiac, and reno-protective activities (Joshi
et al., 2016).

9 Natural Products from Ficus carica for Prevention

of Secondary Complications of Diabetes

• Cardiovascular diseases include diseases/disorders of the heart and blood clots.

Alamgeer et al. (2017) deduced that the presence of phenols, flavonoids, and
potassium contributed to the antihypertensive and cardioinhibitory effects.
However, the beta-adrenergic and calcium channel blocking activity were not
detected, implying the antihypertensive effect was due to some other mecha-
nism. In another study, 70% methanolic extracts of F. carica leaves were used to
prevent ischemia/reperfusion I/R-induced myocardial infarction in isolated rat
hearts (Allahyari et al., 2014).
• Neurodegenerative disorders imply progressive damage to the structure and
function of neurons. It includes diseases like prion disease, Parkinson’s disease,
Alzheimer’s disease, and Huntington’s disease. Since oxidative stress contrib-
utes to the diseases, antioxidants from F. carica, namely hydroxycinnamic acids,
flavonoids (e.g., rutin), etc. (Szwajgier et al., 2017). According to a study, the
n-hexane and acetone extract of the leaves could inhibit activities of the enzyme
acetylcholinesterase (AChE) by 62.9% and 50.8%, and that of butyrylcholines-
terase (BChE) by 76.9% and 45.6%, respectively (Orhan et al., 2011).
• Kidney damage (nephropathy)
Ghaffar et al. (2015) induced nephrotoxicity in adult albino mice using gentami-
cin, thereby increasing their serum urea and creatinine levels. Ficus carica L. leaf
extract was fed orally to a group of mice at a concentration of 400 mg/kg/day
along with gentamicin 200 mg/kg/day intraperitoneally. Their histological struc-
ture was compared with healthy mice and mice with induced nephropathy, and a
marked improvement in the renal structure was observed. In another study,
F. carica supplementation improved the serum ALT (alanine aminotransferase)
and AST (aspartate aminotransferase) levels and ameliorated the metabolic alter-
ations induced by gamma irradiation in male albino rats (Fouad et al., 2019).
F. carica leaves extract was coated with Au nanoparticles to increase the sen-
sitivity and circumvent the potential drug resistance. Au-NPs and extract at the
ratio of 3:2 could significantly decrease blood urea, MDA, homocysteine levels,
serum creatinine, and hydroxyproline content in kidney tissue homogenate of
Cisplatin-induced acute kidney injury (El-Sayed et al., 2019)
• Eye damage (retinopathy)
Güven et al., 2019 extracted fig seed oil and found that the oil was high in
α-linolenic (n-3) acid, one of the Omega 3 fatty acids recommended for ocular
24 Preventive Roles of Phytochemicals from Ficus carica in Diabetes… 553

health. Vitamin E and α-linolenic acids from F. carica are used in supplementary
therapy to treat yellow spot damage or macular degeneration (Richer et al., 2016)
• Foot damage
Most diabetic patients experience peripheral artery disease (PAD), leading to
decreased foot blood flow. If these patients suffer from neuropathy, they lose
sensations in their feet altogether. This subjects the feet to a plethora of problems
ranging from foot ulcers and calluses to increased skin infections, eventually
leading to amputation of the feet (Holt et al., 2021). Phytochemicals from F. car-
ica could prevent neuropathy and prove an essential remedy for diabetes-induced
foot damage.
• Skin conditions
According to American Diabetes Association, skin problems are usually the first
warning signs of diabetes. Bacterial and fungal infections are usually triggered in
diabetic patients. However, several skin conditions are observed chiefly or only
in people with diabetes (Fig. 24.4).
Traditionally, leaf paste and latex of F. carica have been used for treating a wide
array of skin infections. According to Park et al. (2013), ethanolic extract from fig
branches could considerably reduce the production of TNF-α, i.e., a pleiotropic
inflammatory cytokine. In another study, the flavonoids from fig’s skin extract could
inhibit tyrosinase enzyme at IC50 values from 0.09 to 0.45 mg/m: (Meziant et al.,
2021). This would imply it can be used for skin depigmentation observed in skin
conditions resulting from diabetes.

Fig. 24.4 Skin conditions observed in diabetic patients

554 A. Paramanya et al.

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Chapter 25
Composition and Health-Promoting Effects
of Fig (Ficus carica) Extracts

Toyosi Timilehin George, Ayodeji B. Oyenihi, Omolola R. Oyenihi,

and Anthony O. Obilana

1 Introduction

The leading causes of death or morbidity in humans have been projected to be the
development or progression of chronic, non-communicable diseases due to various
consumer lifestyle practices ranging from food consumption habits and environ-
mental and genetic factors. These disorders and life-threatening conditions are
envisaged to be more pronounced among the poor and the most vulnerable societies
and developed countries and societies with unhealthy lifestyles and eating habits.
Much of the curative measures for these disorders widely available are synthetic
drugs, although plant compounds have been reported to be viable sources of impor-
tant healthy compounds against many human diseases (Ajeigbe et al., 2021; Oyenihi
& Smith, 2019). While advances have been made over the years in medicine, phar-
macology, and pharmacy culminating in discovering therapeutic drugs, the side
effects associated with some of these (adverse drug reactions) have been a cause for

T. T. George (*)
Department of Food Science and Technology, Texas A&M University,
College Station, TX, USA
e-mail: [email protected]
A. B. Oyenihi
Functional Foods Research Unit, Faculty of Applied Sciences, Cape Peninsula University
of Technology, Bellville, South Africa
O. R. Oyenihi
Oxidative Stress Research Centre, Department of Biomedical Sciences, Faculty of Health
and Wellness Sciences, Cape Peninsula University of Technology, Bellville, South Africa
A. O. Obilana
Department of Food Science and Technology, Faculty of Applied Sciences, Cape Peninsula
University of Technology, Bellville, South Africa

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 561
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_25
562 T. T. George et al.

concern (Oyenihi & Smith, 2019). This has led to the continuous awareness of the
need for increased consumption of plant-based foods, generally believed to be less
toxic or have well-tolerated safety profiles from centuries of anecdotal usage.
Furthermore, empirical pieces of scientific evidence continue to indicate positive
results in many in vitro and in vivo experimental trials involving these medicinal
plants (Lazreg Aref et al., 2011; Shahinuzzaman et al., 2020; Wojdyło et al., 2016).
These have been ascribed to the plethora of bioactive phyto-compounds with health
benefits against chronic diseases such as obesity, cancer, infertility, and cardiovas-
cular and neurodegenerative disorders. Fig (Ficus carica) is one of such plants that
has been reported to contain rich and healthy compounds that can benefit consumers
as functional food components.
Fig (Ficus carica) tree is one of the earliest cultivated crops in history, belonging
to the Moraceae family (Mulberry family), and originated from the Middle Eastern
region of the world. Because of its health, nutritional and economic importance, the
fruit is now being cultivated in many world areas (Abdel-Rahman et al., 2021;
Wojdyło et al., 2016) and consumed in fresh, dried, and processed forms (Dueñas
et al., 2008). Fig is mainly cultivated by countries such as Turkey, Egypt, Tunisia,
Iran, Morocco, and many other countries on the west and the east Mediterranean.
Countries such as Brazil, the United States, China, and Portugal have increased their
fig production capacity (Veberic & Mikulic-Petkovsek, 2015). The color of the fig
fruit (Ficus carica) varies depending on planting location and sub-species in which
they belong, with purple and green being the most common colors. In addition, the
fig tree has immense cultural and religious significance in areas of the world where
they originated, as they are used as objects of religious and cultural worship
(Badgujar et al., 2014). Beyond their use in various cultural settings, they have good
economic potential due to their desirable sensory and organoleptic properties and
their health-promoting ability. Empirical evidence from scientific research has
detailed the importance of the fig fruits and leaves, especially in human diets and
health, with various parts of the tree reported possessing health-promoting com-
pounds and attributes, potent against many known lifestyle diseases (Abdel-Rahman
et al., 2021; Amessis-Ouchemoukh et al., 2017). To further reinforce this, the
medicinal importance of this tree has previously been demonstrated in the tradi-
tional Ayurveda and Siddha system of medicine common in India and other middle
eastern countries (Badgujar et al., 2014).
Different parts of the fig tree have been studied previously and reported for their
health-promoting importance. For example, the fig seed has been reported to con-
tain bioactive compounds that have antioxidant activities and can mitigate oxygen
imbalances in the body (Nakilcioğlu-Taş & Ötleş, 2021). This may be attributed to
phytosterol and its components, such as β-sitosterol and stigmasterol, which possess
cholesterol-lowering properties (Barolo et al., 2014; Jeong & Lachance, 2001). In
addition to the seed, the fig fruit and leaf have been reported for various therapeutic
importance against chronic and lifestyle diseases such as cancer, diabetes, etc.
(Boyacıoğlu et al., 2021; Wojdyło et al., 2016). For instance, a previous report on
the anticancer property of fig leaf extract shows about 67–80% inhibition of Hep2
and HepG2 cell lines (Abdel-Rahman et al., 2021). In another study, the
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 563

antiproliferative potential of latex from Ficus carica on various HT-29 cancer has
been reported and found promising (Boyacıoğlu et al., 2021).
Generally, the health-promoting function of fig has been attributed to the pres-
ence of bioactive compounds (majorly polyphenols) in different tree parts and may
vary from one part to the other (Kamiloglu & Capanoglu, 2015). Polyphenols are
secondary metabolite compounds present in plants for defense against predators but
possess therapeutic, nutritional, and nutraceutical functions (Veberic & Mikulic-
Petkovsek, 2015). They come in various structural forms and are the largest class of
bioactive compounds in plants hence presenting complexity in their characteriza-
tion (Oyenihi & Smith, 2019). Examples of these compounds include; anthocya-
nins, flavonoids, phenolic acids, and others that have previously been related to the
antioxidant activity of the plant. Polyphenols possess strong chemoprevention and
anti-cancer attributes, and their chemoprevention pathways and mechanisms have
been comprehensively discussed in a study by Oyenihi and Smith (2019). Beyond
their anticancer property, bioactive compounds in figs also possess antioxidants
(Boyacıoğlu et al., 2021; Polat, 2011), antidiabetic (Deepa et al., 2018), and anti-
inflammatory actions.
In this chapter, we focused on the composition and health-promoting attributes
of Ficus carica. We discussed their importance in human health and the economic
benefits of these natural healthy compounds going forward. We briefly mentioned
how the reach bioactive compounds in fig could be preserved and maintained and
recommended the possible inclusion of the health-promoting compounds in fig in
commonly consumed staples.

2 Nutritional and Phytochemical Composition of Fig

2.1 Nutritional Composition of Figs (Ficus carica)

Figs (Ficus carica) are nutritious, containing significant macro- and micro-nutrients
(Al-Snafi, 2017; Chauhan & Tanwar, 2015; Khan et al., 2011; Pal, 2020; Sadia
et al., 2014; Verma et al., 2015; Vora et al., 2017). The quantities and concentration
of these micro and macronutrients depend on differences in varieties, state (dry or
fresh fruit), geographical location, type of soil, and general climate of the fruit’s
environment. The nutritional composition of fresh and dried figs is summarised in
Table 25.1. The dried fruit has been reported to contain a high quantity of carbohy-
drates ranging from 65.2% to 73.5%, while fresh fig fruits contain about 8–20%
(Table 25.1). Most of the carbohydrates in figs are glucose, fructose, and sucrose,
with the balance being dietary fiber, insoluble cellulose in the skin, and soluble
pectin (Pal, 2020). Other proximate compositions of the fruit reported for different
accessions and varieties in different locations are presented in Table 25.1. The nutri-
tional content of the fig is distributed between the peel and the pulp, which are both
edible and have been reported to provide adequate nutritional and biochemical
564 T. T. George et al.

Table 25.1 Nutritional content of fig fruit (g/100 g)

Fresh
Fig fruit fruit Dried fruit References
Moisture 79.9–88.1 12.9–25.9 Al-Snafi (2017), Chauhan & Tanwar (2015), Khan
et al. (2011), Pal (2020), Sadia et al. (2014) and Vora
et al. (2017)
Ash 0.60–4.00 0.89–4.65 Al-Snafi (2017), Chauhan & Tanwar (2015), Khan
et al. (2011), Pal (2020), Sadia et al. (2014) and Vora
et al. (2017)
Protein 0.53–1.30 1.80–8.60 Al-Snafi (2017), Chauhan & Tanwar (2015), Khan
et al. (2011), Pal (2020), Sadia et al. (2014), Verma
et al. (2015) and Vora et al. (2017)
Lipid 0.20–0.53 0.14–2.70 Al-Snafi (2017), Chauhan & Tanwar (2015), Khan
et al. (2011), Pal (2020), Sadia et al. (2014), Verma
et al. (2015) and Vora et al. (2017)
Crude fibre 2.10–2.20 3.68–69.4 Al-Snafi (2017), Khan et al. (2011), Pal (2020), Sadia
et al. (2014) and Vora et al. (2017)
Carbohydrates 7.60–20.0 65.2–73.5 Al-Snafi (2017), Chauhan & Tanwar (2015), Khan
et al. (2011), Pal (2020), Sadia et al. (2014), Verma
et al. (2015) and Vora et al. (2017)
Energy 37.0 308.5– Al-Snafi (2017), Pal (2020), Sadia et al. (2014) and
(Kcal/100 g) 332.7 Verma et al. (2015)

composition required for consumers (Mahmoudi et al., 2018). These authors

reported that fruit pulps were rich in carbohydrates, and the peel of the ‘Onk
Elhamam’, the pulp of the ‘Boughandjo’, and the peel of the ‘Bakkor Khal’ culti-
vars had the highest vitamin C and phosphorus content, as well as rich potassium
and calcium composition.
Figs are rich in various minerals, including Ca, Fe, P, K, and Na, necessary for
the physiological functioning of the human body (Al-Snafi, 2017; Chauhan &
Tanwar, 2015; Mahmoudi et al., 2018; Pal, 2020). As a result of the nutrient content,
Khan et al. (2011) recommended that the fig fruit, which is available all year round,
be used as a natural food supplement to provide instant energy for healthy and
unhealthy persons. This was further buttressed by Vora et al. (2017), who reported
that the concentration of nutrients in dried figs makes them a rich energy source
with a high calorific value and may be consumed as a snack.

2.2 Phytochemical Composition of Figs (Ficus carica)

There are over a hundred bioactive compounds that have been isolated from both
dry and fresh figs (Gibernau et al., 1997; Oliveira et al., 2009; Pal, 2020); some of
these include phenolic acids, flavonoid compounds, anthocyanins, carotenoids, fatty
acids, phytosterols, etc. (Fig. 25.1). The phytochemical composition of figs majorly
depends on the extraction solvent, time, medium, and conditions (Mahmoudi et al.,
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 565

Fig. 25.1 Main phytochemical component of figs (Ficus carica)

2018; Verma et al., 2015). Additionally, different planting conditions, locations, and
varieties may result in variations in phytochemical composition (Çalişkan &
Aytekin, 2011; Veberic et al., 2008). Phytochemical compounds extracted from fig
have been reported to have significant antioxidant activity capable of scavenging
free radicals, thus inhibiting the oxidative mechanisms that lead to degenerative ill-
nesses and life-threatening conditions (Oliveira et al., 2009). Phytochemicals in figs
are distributed in various parts such as peels, pulps, seeds, etc. Mahmoudi et al.
(2018) observed that variable levels of bioactive compounds, including polyphe-
nols, flavonoids, anthocyanins, and tannins, were distributed in the peels and pulps
of some cultivars and concluded that generally, the peels, especially those with a
dark color, contained the high concentrations of phytochemicals and exhibited the
highest antioxidant activity compared to fig fruit pulps. They also recommended the
consumption of the whole fig fruit based on the phytochemical content of the tested
fig cultivars. Shamsi et al. (2020) concluded that figs are a good source of essential
nutrients and bioactive compounds that play a vital role in mitigating the occurrence
of degenerative and non-degenerative diseases and illnesses. Therefore, the con-
sumption and usage of figs as inclusion in daily diets is essential for the proper
functioning of human health.

3 Health-Promoting Effects of Fig

3.1 Antioxidant and Anti-inflammatory

The interconnections between the antioxidant and anti-inflammatory qualities of

natural products, including herbs used as foods or medicines to help prevent or slow
down the development of chronic diseases, have been well described (Serafini &
Peluso, 2016). The biochemical and molecular mechanisms of actions of medicinal
plants commonly utilize the antioxidant and anti-inflammatory pathways to achieve
these therapeutic effects. Assessment of antioxidant activities is usually evaluated
566 T. T. George et al.

Fig. 25.2 Health-promoting effects of fig (Ficus carica)

through the measurement of general antioxidant capacities (e.g., ABTS, FRAP,

DPPH·, ORAC), enzyme activities (e.g., superoxide dismutase (SOD), catalase
(CAT), glutathione peroxidase, etc.) and amount of antioxidant compounds (e.g.,
reduced glutathione (GSH), vitamins C & E, α-lipoic acid, etc.). Additionally, the
expression patterns of antioxidant genes or proteins (notably, Nr-f2 and antioxidant
response elements) are also commonly determined. The ability of fig (Ficus carica)
to influence these antioxidant indices has been well-defined in the literature
(Fig. 25.2). For example, scavenging of DPPH·, ABTS, and hydroxyl superoxide
radicals and inhibition of hydrogen peroxide or xanthine oxidase by fig ethanol
extracts were reported (Amessis-Ouchemoukh et al., 2017; Saoudi & El Feki,
2012). Seasonal variations possibly play a role in the antioxidant potency of fig
leaves, as those collected in July exhibited stronger activity than those from June or
September (Marrelli et al., 2014). The ABTS, FRAP, and DPPH· values also corre-
spond with the polyphenol and carotenoid constituents and indicate a preponder-
ance of antioxidant capacity in the fig peel/skin than pulp or whole fruit (Arvaniti
et al., 2019), although the leaf was initially shown to be more potent than fruit,
wood or bark (Belguith-Hadriche et al., 2017). The antioxidant quality of dark-
colored fig varieties may also be superior to their light-colored counterparts
(Arvaniti et al., 2019; Pereira et al., 2017). This was also demonstrated previously
where the peel and total fruit juices from dark-colored variety had the most DPPH·
and reduced power activities (Harzallah et al., 2016). Furthermore, water extract of
fig leaves was demonstrated to be 3× and 4× higher in ABTS radical scavenging
compared to the ethanol and ultrasound-assisted extracts, respectively (Alcántara
et al., 2020). The flavonoid-rich fig peel extracts were recently shown to exhibit
potent free radical scavenging and metal-chelating activities associated with their
ability to inhibit main enzymes implicated in many chronic diseases such as tyrosi-
nase, 𝛼-glucosidase, urease, and cholinesterase (Meziant et al., 2021). The pre-
treatment with fig stem extract ameliorated the decreases in hepatic SOD, CAT, and
glutathione peroxidase activities in methanol intoxicated rats (Saoudi & El Feki,
2012). A similar result was obtained for the antioxidant enzymes in carbon
tetrachloride-induced hepatotoxicity in mice (Hira et al., 2021). These studies
exemplify the antioxidant attributes of fig.
Inflammatory processes may be initiated by the recruitment of B cells and T
cells; infiltration of macrophages, leukocytes, and monocytes into cells, followed by
the expression of pro-inflammatory chemokines, cytokines, and cell adhesion mol-
ecules (Tasneem et al., 2019). The elevation of pro-inflammatory cytokines and
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 567

chemokines (e.g., IL-1, IL-6, IL-18, C-reactive protein, TNF-α, MCP-1) is accom-
panied by the decrease in those of the anti-inflammatory mediators (e.g., IL-4, IL-10
IL-13, TGF-β) have been the major hallmarks of inflammation-induced cellular
injury (Tasneem et al., 2019). Thus, these markers represent major targets of many
therapeutics and medicinal plants, such as fig. For example, the anti-inflammatory
effects of fig leaf extract have been demonstrated by inhibiting leukocyte accumula-
tion and production of TNF-α, PGE2, or VEGF in a rat air pouch model (Eteraf-
Oskouei et al., 2015). Furthermore, the fig leaf extract was also reported to
downregulate VEGF, TNF-α, and IL-1α gene expressions in human keratinocytes
(Turkoglu et al., 2017). Besides the treatment of immuno-compromised mice with
fermented fig extracts enhanced serum levels of IL-4, IL-6, TNF-α, and IFN-γ, sug-
gesting their immune system-modulatory capacity (Zhao et al., 2020).
Oxidative stress has often correlated with many inflammatory or immune-
modulatory assessments as both phenomena mainly act in concert to propagate
chronic diseases (Ahmed et al., 2017). Many pro-inflammatory processes can be
triggered by excess free radicals or vice versa, thus illustrating the medicinal utility
of fig extracts which could act in both antioxidant and anti-inflammatory mecha-
nisms to achieve efficacy. For instance, the administration of fig seed oil reduced
tissue levels of TNF-α and IL-1β and elevated those of GSH, SOD, and CAT, result-
ing in the amelioration of ischemia and reperfusion injury in rats (Orak et al., 2021).
The silver nanoparticles from fig extracts were also shown to reduce lipid peroxida-
tion and nitric oxide levels and down-regulate the gene expression of TNF-α, and
IL-1β, in a leishmaniasis-induced inflammation mouse model (Almayouf et al.,
2020). In a recent study, the protective effect of fig extract against tebuconazole-
mediated toxicity in bats was attributed to its antioxidant and anti-inflammatory
activities.

3.2 Antidiabetic

Diabetes, a chronic disease with heterogeneous pathogenesis and pathophysiology,

continues to ravage the global population, with adult prevalence expected to reach a
whopping 784 million in 2045 (International Diabetes Federation, 2021). The dis-
ease can be distinguished by the loss of glucose and/or insulin homeostasis regula-
tion in the body. While the administration of insulin has been the mainstay in
managing type 1 diabetes, that of type 2 diabetes requires a combination of diet and
lifestyle modifications, exercise, and the use of glucose-lowering drugs. However
the difficulty in adhering to the daily dosage regimen and issues associated with the
clinical side effects of these drugs have hampered their effectiveness, prompting the
increased global acceptance of medicinal plants with antidiabetic properties
(Oyenihi et al., 2020). In this regard, the traditional use of fig to manage diabetes
and its complications has been reported in folklores (Chawla et al., 2012). This
traditional antidiabetic efficacy has been confirmed in many in vitro and in vivo
568 T. T. George et al.

studies with various postulated mechanisms of action as comprehensively reviewed

elsewhere (Deepa et al., 2018). Fig extracts were demonstrated to inhibit the activi-
ties of key enzymes implicated in the progression of diabetes, such as α-amylase
and α-glucosidase (Mopuri et al., 2018; Wojdyło et al., 2016). In a high-fat diet +
streptozotocin-induced type 2 diabetic rat model, fig extracts ameliorated the altered
activities of metabolizing enzymes, including hepatic glucose-6-phosphatase,
fructose-1,6-bisphosphatase, and hexokinase (Irudayaraj et al., 2017) and restored
the lipid parameters to near-normal values (Arafa et al., 2020). The blood glucose-
lowering properties of fig extracts were also exhibited via the suppression of hepatic
gluconeogenesis in an AMPK-dependent manner (Zhang et al., 2019). Abscisic acid
and ficusin compounds isolated from the fruits and leaves of figs, respectively, have
also shown impressive antidiabetic potentials (Atkinson et al., 2019; Irudayaraj
et al., 2016), with the administration of the former reported to lower the glycemic
and insulinemic indices in healthy adults (Atkinson et al., 2019).

3.3 Anti-cancer Effect

The incidence and prevalence of cancer continue to increase worldwide, posing

immense health, social, and economic burdens to humans, requiring novel and con-
certed efforts to reverse this disturbing trend (Mattiuzzi & Lippi, 2019). Therefore,
medicinal plant extracts and other natural products, either stand-alone or supple-
mentary to the current anticancer management strategies, with varying efficacy
issues and toxicity concerns, have been encouraged (Oyenihi & Smith, 2019). The
traditional anticancer effects of fig and derived compounds have been well-reviewed
previously, especially those relating to their cancer preventative and therapeutic,
antioxidant, and anti-inflammatory qualities (Lansky et al., 2008). Fig extracts dis-
played cytotoxicity against a variety of cancer types, including breast (Zhang et al.,
2018b), prostate, colon (Boyacıoğlu et al., 2021), hepatocellular carcinoma (Abdel-
Rahman et al., 2021; Mustafa et al., 2021), and human laryngeal carcinoma (Abdel-
Rahman et al., 2021). In these studies, the anticancer activities were thought to
occur through the elevation of oxidative stress, apoptosis, cell cycle arrest; decrease
in cell migration; and regulation of molecular signaling. Bergapten and psoralen
from the leaves (Zhang et al., 2018a) and ficin from the latex (Boyacıoğlu et al.,
2021) are some of the important anticancer components in figs. Besides, the silver
nanoparticles from dried fig fruits were reported to retain the cytotoxicity against
the MCF-7 breast cell line (Jacob et al., 2017). The benefits of using fig extracts as
an adjuvant to chemo-photodynamic therapy were also recently demonstrated on
rhabdomyosarcoma cells (Aziz et al., 2021).
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 569

3.4 Anti-diuretic Effects

Diuretic drugs modulate the volume and composition of body fluids, hence used for
treating different clinical conditions involving oedema or volume overloads like
hypertension, heart failure, ascites, nephrotic syndrome, and cirrhosis. Herbal
diuretics are considered safe and cost-effective alternatives to synthetic drugs.
Sruthi et al. (2012) investigated the diuretic activity of ethanolic extract of Ficus
carica whole plant at doses of 200 and 400 mg/kg of body weight in rats based on
parameters such as total urine volume and urine concentration of sodium (Na+),
potassium (K+), and chloride (Cl−). Results showed a significant diuretic effect of
the ethanolic extract treatment based on increased urine volume and urinary excre-
tion of electrolytes; Na+, K+, and Cl-.
Munteanu et al. (2016) also reported the diuretic effects of aqueous and ethanolic
leaf extracts of Ficus carica. The extracts showed a dose-dependent increase in
urine excretion. The aqueous extract (1.4 mg/kg) demonstrated a uricosuric effect
which could benefit patients with elevated blood uric acid levels. The chemical
composition of extracts revealed the presence of flavonoids which may be respon-
sible for inhibiting tubular reabsorption of water and anion and its diuretic activity.
However, it is still not clear if the diuretic activity is attributed solely to the flavo-
noid constituents or the synergistic effects of the phytoconstituents. Therefore, there
is a need to investigate the active principles responsible for the diuretic effects. Also,
these findings agree with the traditional claim of Ficus carica fruits as a diuretic
agent in Unani, Ayurvedic, and Chinese-traditional systems of medicines (Badgujar
et al., 2014; Chawla et al., 2012; Salma et al., 2020). The scientific investigations
regarding the diuretic effects of Ficus carica still require novel in-depth analysis
that can ascertain the needed dosage level for safety.

3.5 Anti-neurodegenerative Effects

Alzheimer’s disease (AD) and other neurodegenerative disorders (NDDs) are char-
acterized by a progressive decline in neuronal function and structure, which eventu-
ally results in neuronal death in the nervous system (Karnati et al., 2015; Koric
et al., 2016; Pugazhenthi et al., 2017, Zhang et al., 2019). The pathological cellular
and molecular events in NDDs include; oxidative stress, neuroinflammation, aber-
rant RNA processing, DNA damage, protein oligomerization and aggregation, axo-
nal transport deficiency, calcium deregulation, impairment in the mitochondrial
function and structure, and abnormal neuron-glia interactions (Dugger & Dickson,
2017). One of the characteristic features of AD is a progressive loss of memory.
Also, the neuropathology of AD is characterized initially by the deposition of senile
plaques composed of amyloid beta-protein (Aβ) and neurofibrillary tangles contain-
ing hyperphosphorylated tau protein brain and later by the loss of neurons and their
processes. Treatment with Ficus carica fruit improved learning and memory in
570 T. T. George et al.

experimental rat models (Sumanth & Mamatha, 2014). Subash et al. (2014) reported
the neuroprotective effect of Ficus carica fruits supplemented diet in a transgenic
(Tg) mouse AD model. Ficus carica treatment improved memory-related behav-
ioral deficits and reduced oxidative damage and plasma Aβ (1–40 and 1–42) levels
in Tg mice.
Cholinergic mechanisms are involved in memory formation, and cholinergic
cells are susceptible to death in AD patients (Maurer & Williams, 2017). Ficus car-
ica fruit and leaf extracts showed potent inhibitory activities against acetylcholines-
terase and butyrylcholinesterase enzymes (Subash et al., 2014; Sumanth &
Mamatha, 2014). The inhibitory action of Ficus carica on these enzymes prolongs
the activity of acetyl- and butyryl-choline neurotransmitters in the synapse
(Breithaupt & Weismann, 2004; Marshall, 2004; Orhan et al., 2011). Also, Aβ pep-
tides could induce Ca2+ influx that leads to increased activity of these enzymes and
Ca2+ mediated oxidative stress (Barbosa et al., 2002). The mitigating effect of figs
against oxidative stress was evident by the inhibition of malondialdehyde accumu-
lation and a reduction in protein carbonyl levels in the cortex and hippocampus of
Tg mice. In addition, fig supplemented-diet prevented a decrease in the antioxidant
enzymes- superoxide dismutase and catalase and attenuated the elevated levels of
glutathione peroxidase and glutathione reductase in brain regions of AD-Tg mice
(Subash et al., 2014). Also, the APPSw2576 transgenic mouse model of AD was
exposed to a Ficus carica fruit-supplemented diet. Essa et al. (2015) reported a
decrease in plasma inflammatory cytokines levels, suppression of Aβ peptide levels
in both cortex and hippocampus, and a significant recovery in brain ATP levels.
Altogether, data from these studies showed the promising therapeutic benefits of
Ficus carica fruits against neurodegenerative diseases such as AD.

3.6 Anti-lipidemic/Cholesterol Limiting Effect

Dyslipidaemia (characterized by abnormal levels of blood lipids) is a high-risk fac-

tor for cardiovascular derangements such as coronary heart diseases, atherosclero-
sis, and myocardial infarction (Hedayatnia et al., 2020). The antihyperlipidemic
effects of Ficus carica leaf extracts are well-reported in rats and humans (Perez
et al., 1999; Serraclara et al., 1998). Ficus carica significantly improved the lipid
profile and decreased adipogenic risk factors in high-fat diet (HFD) rats (Joerin
et al., 2014). HFD-induced obese male rats fed with Ficus carica leaf extract had
significantly decreased triglycerides and elevated high-density lipoprotein choles-
terol (HDL-C). Another study reported that hydro-methanolic leaf extracts of Ficus
carica and its fractions significantly lowered cholesterol levels in the serum and
liver of experimentally-induced nutritional hyperlipidemic rats (Rassouli
et al., 2010).
Studies also suggest that figs treatment reduced cholesterol levels in an obesity-
linked type 2 diabetes mellitus induced with a high-fat diet (HFD) plus streptozocin
(Arafa et al., 2020). Hydroethanolic extract of the fruit of Ficus carica administered
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 571

at 250 mg/kg/day and 500 mg/kg/day lowered cholesterol, triglycerides, low-density

lipoprotein (LDL), and very-low-density lipoprotein (VLDL) while increasing the
protective effects of high-density lipoprotein (HDL). Furthermore, ethanolic fruit
extract of Ficus carica inhibited pancreatic lipase- an enzyme that digests dietary
fats and enables absorption into the intestinal lumen (Birari & Bhutari, 2007).
Altogether, these studies indicate that Ficus carica protects against cholesterol and
triglycerides accumulation and has the potential to treat or manage atherosclerosis
and coronary heart disorders. These reports also support the traditional application
and benefits of Ficus carica in treating metabolic syndrome-related conditions
(Mawa et al., 2013). The lipid-lowering effects of Ficus carica may be due to the
flavonoids and tannins that play a significant role in the mobilization and metabo-
lism of lipids (Rassouli et al., 2010). Future studies evaluating the mechanisms by
which Ficus carica mediates its antihyperlipidemic activities would provide addi-
tional relevant information.

3.7 Hepatoprotective Effect

Figs have exerted organ-level protection, in the liver in particular, from various inju-
ries caused by chemicals such as ethanol (Turan & Celik, 2016) and carbon tetra-
chloride (Aghel et al., 2010; Singab et al., 2010), and rifampicin (Gond & Khadabadi,
2008). The hepatoprotective effects were suggested to occur via the amelioration of
hepatic marker enzymes (mostly alanine and aspartate transaminases), antioxidant,
and anti-inflammatory actions. In a gamma-radiation toxicity rat model, the daily
administration of fig extract for 21 days elicited decreases in the extent of hepatic
and renal damage, probably through its antioxidant and anti-apoptotic actions
(Fouad et al., 2019).

3.8 Anti-hypertensive Effect

In many parts of the world, most medications available for hypertension and cardio-
vascular diseases are synthetic drugs; however, some have exhibited side effects.
For example, the use of some anti-hypertensive drugs has been linked to cases of
erectile dysfunction (Ajeigbe et al., 2021). The use of medicinal plants and fruits to
treat this life-threatening disease is currently being explored, and the fig is one of
such fruits. This has been linked to the rich presence of antioxidant compounds
present in its fruits, leaves, and seeds (Badgujar et al., 2014; Olomon et al., 2006).
A recent study by Alamgeer et al. (2017) has shown the effectiveness of methanol
extract of fig fruit in decreasing hypertension in normotensive and glucose-induced
hypertensive rats. This study showed that this effect is dose-dependent, and the
effective dosage level was reported to be 1000 mg/kg body weight. The study attrib-
uted this effectiveness of the fig extract to the rich presence of phenolic acids,
572 T. T. George et al.

flavonoids, and potassium in the fruit. This indicates the promising potential of fig
in treating one of the world’s leading causes of death. Since different parts of the fig
tree have been established to contain rich antioxidant compounds, the leaves, seeds,
and barks should be further explored in treating hypertension. A comparison of the
efficacy of the extracts obtained from these different plant parts will also assist in
the isolation and elucidation of the specific compound(s) most responsible for the
anti-hypertensive activity. The paucity of clinical trials on the effectiveness of the
fig extracts in treating hypertension and related health disorders is observed, limit-
ing their clinical usage.

4 Storage and Preservation of Bioactive Contents

of Ficus carica

The shelf life of most fruits and leaves of importance is low primarily due to con-
tinuous respiration during storage and the presence of ethylene (Kaur et al., 2021).
Some of these fruits are rich in bioactive compounds that are essential for promot-
ing consumers’ health. Hence, they must be adequately stored to maximize their
importance. Figs are commonly preserved by freezing, storage in modified atmo-
sphere packages, dehydration, processing into puree for complete freezing, and,
more recently, encapsulating these bioactive contents into nano- and micro-capsules.
These methods have been stated to be a viable preservatives for bioactive plants
(Kaur et al., 2021). Having a system in place that caters, maintains, and preserves
the integrity of healthy compounds in figs will improve its economic value and the
use of the crop in various parts of the world, especially in areas where they are
not native.

5 Conclusion

Although figs have been reported to contain a high amount of essential phytochemi-
cals with health-promoting functions, the medical benefits of the crop are still
largely untapped, especially in areas of the world where they are not native. With
continuous growth in many life-threatening diseases, it is expedient that more medi-
cal and pharmacological research studies are conducted using this nutrient-rich and
health-benefiting plant either via direct supplementation or inclusion in different
human foods. This inclusion can be achieved by fortifying and enriching commonly
consumed staple foods with figs. Hence, research should be placed on determining
the optimal dose required for an effective or efficient fortification protocol.
Additionally, the medicinal benefits of figs should be confirmed in comprehensive,
well-designed clinical studies that include clinical end-points and therapeutic dose-
response and toxicity profile assessments. Also, since most phytochemical-rich
25 Composition and Health-Promoting Effects of Fig (Ficus carica) Extracts 573

fruits are prone to faster degradation due to increased respiration and ethelyne pro-
duction, it is important to find sustainable, cost-friendly, and safe storage and pres-
ervation of figs beyond the current drying technique is frequently employed. This is
important in making them available all year round with their bioactive and phyto-
chemical contents kept intact.

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Chapter 26
Genotoxic and Antimutagenic Activity
of Ficus carica Extracts

Nusrath Yasmeen, Gondrala Usha kiranmai, and Aga Syed Sameer

Abbreviations

NER Nucleotide excision repair

BER base excision repair
CA chromosomal Aberration
SCE sister chromatid exchange
UDS unscheduled DNA synthesis
MN Micronucleus test
HPRT Hypoxanthine-guanine phosphoribosyl transferase
XPRT xanthine guanine phosphoribosyl transferase
MI Mitotic Index
ERA E. coli WP2 reversion assay
SMART Somatic Mutations and Recombination Test
AC Allium cepa Assay
RXL Ring-X-Loss test
SHA Sperm Head Anomaly
AI Apoptosis-Inducing

N. Yasmeen
Department of Biotechnology, Amity University, Jaipur, Rajasthan, India
G. Usha kiranmai
Faculty of Pharmaceutics, University College of Pharmaceutical Sciences, Kakatiya
University, Warangal, Telangana, India
A. S. Sameer (*)
Department of Basic Medical Sciences, Quality Assurance Unit, College of Medicine, King
Saud Bin Abdul Aziz University for Health Sciences (KSAU-HS), King Abdullah
International Medical Research Centre (KAIMRC), Jeddah, Saudi Arabia
National Guard Health Affairs (NGHA), King Abdulaziz Medical City, King Abdullah
International Medical Research Centre (KAIMRC), Jeddah, Saudi Arabia
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 579
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_26
580 N. Yasmeen et al.

RI Cell Replication Indice

PDB Plasmid DNA Breakage
FM Forward Mutagenesis
ICPM Induction of Cytoplasmic Petite Mutations
CYP450 cytochrome P450

1 Introduction

Plant phytochemicals and their secondary metabolites have a tremendous potential

to treat numerous diseases. They are considered safer and economical hence relied
upon by many people globally (Gorlenko et al., 2020). According to the World
Health Organization (WHO), people using traditional medicine to cure their dis-
eases vary between 70% and 80% of the global population. Numerous drugs we are
currently using are derived from natural products, i.e., folk remedies, semi-synthetic
or synthetic (Oyebode et al., 2016). Despite rapid technological advancements and
novel methodologies involved in the production of pharmaceuticals, plant phyto-
chemicals remain an essential source of lead molecular entities for the development
of new drugs. This could be attributed to the fact that phytochemicals are complex
and challenging to synthesize by chemical processes (Forni et al., 2019; Thomford
et al., 2018). Since time immemorial, drugs from plants and animals have been used
based on trial and error methods that exhibited specific therapeutic properties signi-
fied by the cause-and effect-relationship (Atanasov et al., 2015). Traditional medi-
cines used were complex concoctions of naturally occurring substances of varied
origins (plants, animals, mineral sources), etc., whereas modern pharmaceutical
drugs are substances synthetic in origin with a single active entity (Mushtaq et al.,
2018). Existing literature supports that the drugs from crude natural products are
less toxic than single entity synthetic drugs; this may be attributed to several syner-
gistic/antagonistic components in the concoctions.
The biological significance of herbal medicine has led to the resurgence of scien-
tific interest among ethno pharmacologists striving to develop new drugs using
active phytochemical ingredients as a lead molecule. Furthermore, the lack of a
strong regulatory system involved in evaluating, testing, monitoring, and regulariz-
ing natural products as medicines led to the popularity of natural products. The
people’s acceptance of traditional or folklore medicine is based on the belief that
natural products are much safer than their alternative allopathic forms. However,
current scientific data contrasts this general perception and suggests that traditional
herbal medicines can be exceptionally harmful to humans; they can potentially
interact with food and other ingredients and lead to adverse events (Fokunang et al.,
2020; Ekor, 2014). Natural product safety profiling is further jeopardized by inap-
propriate manufacturing, labeling, deceptive claims, and misleading patients.
Therefore, safety evaluation and regulatory policies must be standardized, and strin-
gent regulatory policies regarding herbal medicine usage must be implemented
globally (Alostad et al., 2020).
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 581

The tremendous therapeutic potential of herbal medicine and the belief that
“Natural means safe” clubbed with their long-term usage has led to a lack of signifi-
cant scientific evidence related to herbal drug safety and toxicity parameters.
However, current data and reports about the toxicity effects of herbal medicines
show a significant upraise in acute and chronic intoxications. Additionally, the
availability of antidotes to treat such intoxications and the toxicological impact of
herbal medicine among pediatrics and insensitive groups is unavailable.
Subsequently, it becomes necessary to evaluate herbal medicine (Ghorani-Azam
et al., 2018). Therefore, traditional or folklore medicine must undergo rigorous
screening procedures via several in vitro and in vivo experiments regarding their
safety, efficiency, and toxicity, similar to allopathic drugs.
Furthermore, the preclinical evaluation of herbal medicine must be performed to
verify its mutagenic, genotoxic, and cytotoxic effects (Moreira et al., 2014).
Assessment of the genotoxicity effects of natural medicine is crucial as genotoxins,
if present in phytochemicals, may cause alteration in DNA via mutations and
increase the risk of disease progression of pathologies such as cancer (Qari et al.,
2021). Additionally, toxicological risk assessment can prove beneficial in identify-
ing plant products that possess antimutagenic, anticarcinogenic effects, which can
be considered crucial in the research for developing novel therapeutic agents.
Furthermore, as suggested by Graz (2013), reverse pharmacology, also known as
the “bedside-to-bench” approach, might enhance the knowledge of the safety of
folklore medicines. This chapter summarizes the toxicity risk assessment regarding
genotoxicity and antimutagenic effects of Ficus species, focusing on Ficus carica.

2 Genotoxicity and Its Assessment Techniques

Genetic toxicology (genotoxicity) is a kind of toxicity that describes the ability of

chemical substances to induce changes in DNA or chromosome, enzymes, or cel-
lular organelles of somatic or germ cells, either through direct or indirect interac-
tions, that cause damage/harm to genetic information, but does not necessarily
cause a mutation (Hasselgren et al., 2019). Genotoxicity is involved in specific cor-
rection procedures like the direct repair mutations utilizing nucleotide excision
repair (NER) and base excision repair (BER), or they may exist in somatic cells or
remain as an altered gene in germ cells and cause genetic diseases (Qari et al.,
2021). At the same time, mutagenicity refers to the induction of variations in the
genetic material of organisms that are permanent and transmissible by chemicals
called mutagens. Mutagenic chemicals interact with DNA resulting in alteration in
DNA base pair sequence via addition, deletion, or replacement of bases, thus alter-
ing their sequence in the DNA and conformity of the genetic information, conse-
quently enhancing the mutation frequency, loss of heterozygosity, chromosomal
aberrations (Honma, 2020). These events have a crucial role in various malignan-
cies, including cancer. Therefore, identifying genotoxic, mutagenic, and carcino-
genic effects of traditional medicine, dietary substances, pharmaceuticals, actually
582 N. Yasmeen et al.

every new compound is essential for safety concerns. Plants that possess mutagenic
properties are regarded as potentially unsafe. Hence, toxicity assessment must be
performed as per several regulatory authorities’ terms, and the compounds must be
deemed safe for human use only if these toxicity parameters are ruled out (Wiesner,
2014). To assess genotoxicity and/or mutagenicity, one single endpoint does not
exist, it involves an array of endpoints. The three crucial endpoints which must be
evaluated for genotoxicity assessment are Gene mutation (mutagenicity) frequency,
chromosomal aberration in structure (Clastogenicity), and chromosomal aberration
in number (Aneugenicity). A battery of the most common in vivo and in vitro testing
techniques for genotoxicity utilized are the bacterial Salmonella mutagenicity test
(Ames test), the VITOTOX Assay (VIT), alkaline comet assay, chromosomal
Aberration (CA) assay, sister chromatid exchange (SCE) assay, unscheduled DNA
synthesis (UDS) test, Micronucleus test (MN), Hypoxanthine-guanine phosphori-
bosyl transferase (HPRT) test, a transgene of xanthine guanine phosphoribosyl
transferase (XPRT) test, Assessment of Mitotic Index (MI), E. coli WP2 reversion
assay (ERA), Somatic Mutations and Recombination Test (SMART), Allium cepa
Assay (AC), Umu-C Assay(UC), SOS Induc test(SOS), Assessment of Somatic
Segregation in Aspergillus nidulans (AN), Gene Mutation in Saccharomyces cere-
visiae (SC), Assessment of Recessive Mutation Lethal X-linked (SLRL) in
Drosophilla melanogaster, Ring-X-Loss test (RXL), Sperm Head Anomaly (SHA),
Apoptosis-Inducing (AI), Cell Replication Indice (RI), Plasmid DNA Breakage
(PDB), Forward Mutagenesis (FM), and Induction of Cytoplasmic Petite Mutations
(ICPM) (Sponchiado et al., 2016; Hasselgren et al., 2019; Turkez et al., 2017). Most
commonly used genotoxicity assays are shown in Fig. 26.1.
Conversely, plants with antimutagenic potential are therapeutically significant
due to their chemo-preventive properties (Ntuli et al., 2018). Chemical agents that
reduce the mutagenicity of chemical or physical substances, either by inhibiting the

Fig. 26.1 Commonly used genetic toxicity tests (in vitro and in vivo)
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 583

action between DNA and mutagens or by neutralizing the mutagen itself, are known
as Antimutagens. The term antimutagens were coined by Novick and Szilard (1952).
The antimutagenic agents were classified as desmutagens and bioantimutagens.
Desmutagens are agents that suppress mutation by blocking the chemical interac-
tions, whereas, Bioantimutagens act intracellularly after the DNA damage occurs,
processing its “repair and replication”, subsequently decreasing the mutation fre-
quency (Choudhuri et al., 2021; Dashwood, 2002). Natural products exhibit their
antimutagenic properties via any one of the following mechanisms (1) Inactivation/
sequestration of mutagens directly, (2) enhanced deactivation or inhibition of meta-
bolic activation of mutagen, (3) protection of DNA against toxic effects of muta-
gens, (4) Rectification of DNA damage via repair and replication to make it less
mutagenic, (5) cell cycle modulation, (6) interference with detoxification system of
the liver (Akram et al., 2020; Zeiger, 2011). The investigation of antigenotoxic natu-
ral products must be done to develop a novel anti-cancer drug, as severe side effects
and drug resistance accompany the current therapeutic strategies for cancer. Hence,
replacement with a less hazardous and more beneficial product is anticipated.
Interestingly, certain compounds possess both mutagenic and antimutagenic
effects. These compounds are known as “Janus mutagens” (Słoczyńska et al., 2014).
For instance, the lignin dibenzyl butyrolactone called hinokinin (HK) obtained from
the dry seeds of Piper cubeba act as a “janus” compound, i.e., at low doses, behaves
as an antioxidant/antimutagen (scavenge ROS). In contrast, at high doses, it is a pro-
oxidant/mutagen (which produces free radicals) (Resende et al., 2012).
Unfortunately, these biphasic properties of phytochemical compounds are not tested
appropriately for their mutagenicity/genotoxicity/carcinogenicity, yet they are rec-
ommended as potent anticarcinogens for human use. This warrants a thorough
screening of natural products before being used as medicines, and the screening
reports must be interpreted with extreme care and caution (Errol Zeiger, 2003).

3 Overview of Ficus carica

Ficus is considered the largest genus of angiosperms, with approximately 850 spe-
cies identified and distributed worldwide, covering tropical and sub-tropical zones
(Yang et al., 2015). Plants belonging to this genus are deciduous shrubs or small
evergreen trees, herbs, creepers or climbers, epiphytes or hemi epiphytes, litho-
phytes, and vines rheophytes (Suttisansanee et al., 2021). Plants under the genus
Ficus belong to the order Urticales and the family Moraceae (Mulberry). They have
nurtured crucial niches in economic and therapeutic areas. Numerous Ficus spp.,
with their parts like stem, roots, bark, leaves, fruits, latex, resins, etc., are deemed
medicinally important in Ayurveda, Chinese, and Arabic folklore medicine as sev-
eral pharmacological activities have been reported (Irudayaraj et al., 2017). Among
the several Ficus species, Ficus carica Linn. (FC) has gained significant popularity
due to its edible fruit, either in dry or fresh form, either peeled or unpeeled, and is
commonly known as “fig”. It is the oldest known cultivated fruit native to the
584 N. Yasmeen et al.

Mediterranean Basin, the Middle East, and southwest Asia. F. carica grows up to a
height of 6.9–10 m, has a small grey bark and numerous branches which secrete
milky white latex, enriched with enzyme Ficin (Hamed et al., 2020), and fragrant
leaves that are oval, orbicular in shape, the pubescent undersurface and above the
rough surface with 3–5 lobes (Patil & Patil, 2011). The inflorescence of Ficus carica
is complex and consists of Synconium with unisexual flowers (Peniwidiyanti et al.,
2021). However, fruits are mature synconia, fleshy, hollow, and involuted with a
closed receptacle. They are bulbus with ostiole (opening) at the distal end for pol-
lination by female wasp Blastophaga psenes (L.) Grav. Their peel color varies as it
ripens from green to purple, attributed to the presence of anthocyanins (Teixeira
et al., 2018; Wang et al., 2017). In recent times figs have gained enormous attention
as they constitute an integral part of the Mediterranean diet. Fig is a highly nutri-
tious fruit, low in sodium, abundant in vitamins (A, B1, B2, B3, and C), amino acids
(aspartic acid and glutamine), minerals (potassium, calcium, magnesium, zinc,
iron), natural sugars; they have no cholesterol or fat, polypeptides and phenolic
compounds (Mohammed et al., 2020; Pereira et al., 2017).
Ficus carica biosynthesizes several primary and secondary metabolites that can
be isolated from its plant parts such as leaves, bark, dried and fresh fruit, i.e., fig,
latex. Approximately 126 chemical constituents have been identified in F. carica to
date, many more are yet to be revealed. The major type of phytochemicals reported
in Ficus carica is phenolic acids, flavonoids (mostly in fruits and leaves), carot-
enoids, anthocyanins (fruit pulp and peel), coumarins (leaves and roots),
sterols(leaves), and terpenoids, alkaloids, tannins, sterols and proteins (latex), arabi-
nose, β-carotenes, β-amyrins, glycosides, xanthotoxol, and β-sitosterol. Arvaniti
et al., in their study dealing with fresh and dried figs, reported the presence of gallic
acid, rutin, chlorogenic acid, epicatechin, and quercetin-3-O-rutinoside (Arvaniti
et al., 2019). Anthocyanins of type, pelargonidin-3-O-rutinoside, cyanidin-3,5-
diglucoside, and cyanidin-3-O-rutinoside were predominantly present in fig peel,
whereas epicatechin and cyanidin-3-O-rutinoside were present in pulp extracts
(Hssaini et al., 2021). Various phytochemicals of therapeutic significance identified
in plant parts of Ficus carica like root, fruit, leaves, latex, etc., are enlisted in
Table 26.1, while their chemical structure is depicted in Fig. 26.2.

4 Therapeutic Potential of Ficus carica

Various parts of Ficus carica like leaves, fruit, latex, and root are deemed medici-
nally valuable in folklore medicine and traditionally used to treat ailments. Ficus
carica treats diarrhea, expectorant, diuretic, colic, intestinal spasm, and gastrointes-
tinal tract indigestion. Similarly, respiratory disorders such as sore throats, bron-
chial problems and inflammation, and cardiovascular and metabolic disorders are
treated (Baygeldi et al., 2021). The utilization of “fig” fruit as food and medicine is
as old as the human civilization. In the ancient Indian medicine system called
Ayurveda, sore throat was relieved by a gargle of fig decoction, also used to treat
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 585

Table 26.1 Phytochemical constituents of therapeutic significance identified in various parts

(roots, leaves, fig, latex) of Ficus carica
Part used Class of phytochemical constituent Example
Leaf Coumarin 4′,5′-Dihydropsoralen
Umbelliferone
Marmesin
Bergapten
Flavonoid Rutin
Sterol Baurenol
24-methylenecycloartanol
Lupeol
ψ-taraxasterol ester
Triterpenoid Ficusogenin
Leaf, root Coumarin Psoralen
Sterol β-Sitosterol
Fruit Coumarin Umbelliferone
Scopoletin
Alkaloid Quinines
Anthocyanin Cyanidin-3-O-glucoside
Cyanidin-3- O-rhamnoglucoside
Hydrocarbon Stilbenes
Latex Triterpenoid 6-O-Linoleyl-b-D-glucosyl-b-sitosterol
6-O-Oleyl-b-D-glucosyl-b-sitosterol
6-O-palmitoyl-b-D-glucosyl-b-sitosterol
Sterols Betulol
Lupeol
Lanosterol
Lupeol acetate
β-amyrin
β-sitosterol
α-amyrin
Acyl steryl glucosides 24-Methylenecycloartanol
ψ-taraxasterol ester

dysentery, menorrhagia, and gonorrhea, and used as an aphrodisiac. Not only this,
Ayurveda recommends ingestion of entire fig for treatment of diarrhea. Evidence
from traditional Chinese medicine recommends a decoction of fig for local applica-
tion and cooked fig for ingestion to cure hemorrhoids. Traditionally, the decoction
of the Ficus leaves was used to treat diabetes (Konyalιoğlu et al., 2005). Italian
people used a decoction of dried leaves along with apple peel as a cough suppres-
sant; this decoction can have walnut hulls, leaves of Malva sylvestris, and heads of
Matricaria chamomilla (Mautone et al., 2019).
F. carica latex was applied to treat warts and viruses. This traditional use was
substantiated by the evidence provided by Bohlooli et al., who reported no adverse
effects and safety in the use of folklore medicine (Bohlooli et al., 2007). Additionally,
586 N. Yasmeen et al.

Fig. 26.2 Chemical structures of phytochemical constituents of therapeutic significance in Ficus

carica. (Structures were obtained from https://pubchem.ncbi.nlm.nih.gov/compound/; http://www.
chemspider.com/Chemical-Structure.XX.html)

dried figs soaked in olive oil were a traditional cure for cough and bronchial prob-
lems. Manjula et al. reported that jaundice could be treated when 20 mL of fig leaf
juice is administered with a cup of goat milk once a day for 3 days (Manjula et al.,
2011). The green unripe fig fruits, when cooked, can serve as galactogogue and
tonic; upon roasting, the fruit acts as an emollient and a poultice that can be used to
treat gumboils, dental abscesses (Ali et al., 2009). Fig is a folkloric emmenagogue
used to treat amenorrhea (Salehi et al., 2021). Although traditional phytotherapy is
based on experience and is transferred to generations by word of mouth, very mea-
ger evidence exists in the form of documents. Undoubtedly, fig has many more uses
which need to be investigated. Based on the available folkloric use, scientists have
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 587

conducted experiments and tried to prove the uses of Ficus carica. Consequently, it
was discovered that Ficus carica L. (fig) possesses several pharmacological proper-
ties such as antispasmodic, anti-cancer, anti-inflammatory, antioxidant, antiviral,
hemostatic, etc. (Mawa et al., 2013). The most prominent therapeutic properties of
fig are summarized in Table 26.2. Although the exact mechanisms of action of Ficus
carica used for the treatment of various disorders are not fully elucidated, it is
apparent that the ubiquity of polyphenols, high content of total phenols, and flavo-
noids are responsible for the actions. The therapeutically significant phytoconstitu-
ents of fig might exert their action through any of the following mechanisms: radical
scavenging antioxidant effects, antiproliferative effects, apoptotic induction, induc-
ing cytochrome P450 (CYP450) monooxygenase enzyme system, which ultimately
enhances the body’s innate detoxification system. About its varied uses, Ficus car-
ica has occupied ethnopharmacological niches and has garnered a place in occiden-
tal Pharmacopeias, in PDR for Herbal Medicines, and is a compendium of herbology.
As one of the oldest cultivated crops, Ficus carica fruits are considered safe.
However, specific toxicity issues have been reported related to FC leaves, as to
moderate toxicity and causation of phototoxicity. Furthermore, exposure to latex
may initiate inflammatory reactions like asthma, dermatitis, asthma, and/or anaphy-
laxis. Furthermore, this oral administration of latex may initiate hallucinations
(Lansky et al., 2008). Hence evidence must be gathered based on scientific investi-
gations, and the entire toxicity profile must be updated; meanwhile, FC leaves must
be consumed with precaution. This is possible only by a series of toxicological
investigations.

Table 26.2 The most prominent therapeutic properties of Ficus carica

Therapeutic
property Plant part used Mechanism Reference
Anticonvulsant Stem bark ethanolic extract Potentiation of the Raafat and Wurglics
activity inhibitory glycine (2019)
receptor
Hepatoprotective Petroleum ether extract of Presence of flavonoids; Gond and
activity leaves; ethanolic leaf detailed mechanism yet Khadabadi (2008),
extract; methanolic leaf to be known Mujeeb et al. (2011),
extract; 80%aqueous Mohan et al. (2007),
ethanolic leaf extract and Aghel et al.
(2011)
Antioxidant Methanolic leaves extract Free radical scavenging Ayoub et al. (2019)
activity free-radical scavenging potential and Ali et al. (2012)
effect, hydroalcoholic
extract of leaves
Anti- Hydro-alcoholic extract of Scavenge the free Ali et al. (2012) and
inflammatory F. carica leaves; ethanolic radicals like superoxide Park et al. (2013)
activity extract and fractionation of generated by
fig branches photoreduction of
riboflavin; inhibit NO
and TNF-α in
RAW264.7 cells
(continued)
588 N. Yasmeen et al.

Table 26.2 (continued)

Therapeutic
property Plant part used Mechanism Reference
Alzheimer’s Hexane leaf extract; a AChE inhibition; Saxena et al. (2013),
disease mixture of fig and olive oil; phenolic and flavonoid- Alharthy and
dietary supplementation yet unclear Bawazir (2019), and
phenolic compounds and Subash et al. (2016)
flavonoids
Anti-diabetic Methanolic extract of stem The insulinotropic effect Bhat et al. (2013),
activity bark; ethanolic fruit of the extract; inhibition Mopuri and Islam
extract; freeze-dried leaf of α-amylase and (2016), and Zhang
extract α-glucosidase enzymes; et al. (2020)
inhibits apoptosis in
pancreatic β-cell via
inhibition of AMPK/
JNK/caspase-3 pathway
Hypoglycaemic Ethyl acetate extract of Insulin sensitization Stephen Irudayaraj
activity leaves; aqueous leaf extract activity; prevention of et al. (2017) and
B-cell degeneration; Perez et al. (2000)
enhanced insulin
secretion
Anticancer Methanolic leave extract; Preventing cellular Sadeghizade et al.
activity methanolic leaf and fruit proliferation, enhancing (2021), Purnamasari
extract on Huh7it liver apoptosis in melanoma et al. (2019), Soltana
cancer cells; Peel, pulp, cancer cells; luteolin et al. (2019),
leaves, whole fruit, and and quercetin induce Hashemi et al.
latex extracts on HT-29 cell cycle arrest and (2011), Jacob et al.
and HCT-116 colorectal enhance apoptosis by (2017), Ghanbari
cancer cells; fig tree latex; activating p53 and p21; et al. (2019), and
F. carica fruit latex; apoptosis inducers; AlGhalban et al.
synthesized silver antiproliferative; (2021)
nanoparticles of dried fig; downregulation of p16
latex; and HPV oncoproteins;
cytotoxic effect on
MCF7 breast cancer
Cell lines; downregulate
ERK2, CREB, and
AKT2 pathways in
MDA-Mb-231 breast
cancer cells
Antibacterial and Methanolic extracts of Not clear Shafique et al.
antifungal activity fruit, leaf, root, bark (2021)
Nematicidal Methanolic leaf extract Presence of psoralen Liu et al. (2011)
activity
Antispasmodic 80% ethanolic dried fruit K+ ATP channel Gilani et al. (2008)
and antiplatelet extract activation
activity
(continued)
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 589

Table 26.2 (continued)

Therapeutic
property Plant part used Mechanism Reference
Hemostatic effect latex Ficin shows a Richter et al. (2002)
haemostatic effect
through activation of
Factor X
Anti-nociceptive Petroleum ether extract of Yet to be elucidated Mirghazanfari et al.
effect delayed fruits (2019)
effect
Antihypertensive Aqueous-methanol extract Polyphenols leading to Alamgeer et al.
and of fruits vasorelaxation might be (2017)
cardioinhibitory involved in
activity antihypertensive effects
yet to be elucidated

5 Genotoxicity and Antimutagenic Evidence for Ficus carica

The global trend is to use natural phytochemicals in food and medicine. Ficus car-
ica (Fig) products can be considered the best examples of natural products used in
food and medicine. Ficus carica consists of several phytoconstituents of therapeutic
value. Despite being deemed safe since its usage from ancient times, it can also
exhibit toxicity effects. Phytoconstituents can be genoprotective if they offer protec-
tion against DNA damage, cause inactivation of carcinogens, or enhance DNA
repair processes. Genoprotective/antimtagenic compounds can be anticarcinogenic/
chemopreventive. Hence, it is becoming imperative to investigate compounds with
the above-mentioned activities, so they can be used to develop new drugs.
The anti-mutagenic potential of figs is not thoroughly studied, and the research
towards this toxicity assessment is still naïve. The first antimutagenic action of
Ficus carica extract was on environment xenobiotics reported. Agabeili and kasi-
imova observed a decrease in mutation frequency induced by environmental muta-
gens such as N-metil-N′-nitro-N-nitrozoguanidin (MNNG) when treated with
extracts of Armoracia rusticana, Ficus carica, Zea mays, and their mixture decreased
the level of mutations in Vicia faba cells, chlorophyll mutations in Arabidopsis
thaliana, and NaF-induced mutations in rat marrow cells (Agabeili & Kasimova,
2005). Furthermore, Radwan et al. (2018) investigated the genotoxic effects of
Ficus carica and Schinus molle in varied climatic conditions (i.e., in healthy and
polluted) in the Asir region of Saudi Arabia. They carried out Genotoxicity tests in
mice using Comet assay, micronucleus test, and chromosomal analysis. They con-
cluded from a comet assay that F. carica and S. molle collected from polluted areas
significantly damaged the DNA in bone marrow cells and increased the number of
micro-nucleated cells in the mice. Briefly, plants from polluted areas are genotoxic
to bone marrow cells in mice. They also deduced that metals like Ni and Co are
essential for certain cellular processes. However, their bioaccumulation leads to
chromosomal aberrations, breaks in single or double DNA strands, and a cross-link
between chromatid exchange DNA-protein or DNA damage (Radwan et al., 2018).
590 N. Yasmeen et al.

Cisplatin is a platinum-based antineoplastic agent successfully employed to treat

several cancer types like breast cancer, ovarian cancer, neuroblastoma, brain tumors,
etc. The main disadvantage of current cancer chemotherapies is resistance and toxic
side effects. Cisplatin also subjects the patients using it to nephrotoxicity, genotox-
icity, Hepatotoxicity, and reproductive toxicity (Makovec, 2019). Fahmy et al. eval-
uated the genoprotective/antimutagenic effect of Ficus carica (fig) utilizing
methanolic leaf extracts against cisplatin-induced genotoxicity and testicular dam-
age. Genetic toxicology endpoints worked on here are bone marrow micronuclei in
polychromatic erythrocytes (MPEs), chromosomal aberrations (CAs) in the bone
marrow, and primary spermatocytes with the expression of the genes TNF-α, iNOS
and NF-kB. Cisplatin treatment caused a significant increase in the percentage of
chromosomal abnormalities, increased gene NF-kB levels, increased micronuclei in
polychromatic erythrocytes, and enhanced toxicity to bone marrow cells. However,
treatment with Ficus carica extract reversed all the abnormalities in a dose-
dependent manner proving it to be a promiscuous chemopreventive/genoprotective
agent (Fahmy et al., 2020).
Lightbourn and Thomas further investigated the protective effect of ficus leaf
extract against DES-induced DNA damage in MCF10A cells which are benign
human breast epithelial cells also they investigated the nuclear insult induced by
environmental estrogens and their metabolites on MCF10A cells using Single-cell
gel electrophoresis (SCGE)/COMET assay. Diethylstilbestrol (DES) is a carcino-
genic agent. It is a synthetic estrogenic hormone that is acted upon by CYP450
enzymes converting it to an intermediary metabolite DES-4,4′-quinone (DESQ)
which forms DNA adducts and strand breaks.

Cytochrome p450
Diethylstilbestrol DES-4,4’-quinone DNA adducts and
strand breaks
(DES) (DESQ)

DNA Damage

In this study, they concluded that Ficus carica L. leaf extract inhibits DNA strand
breaks by interfering with the nuclear complex in MCF10A cells either in the pres-
ence/absence of diethylstilbestrol (DES) and 4′,4″-diethylstilbestrol quinine
(DESQ) metabolite (Lightbourn & Thomas, 2019).
Hadžic et al. studied the inhibitory effects of delphinidin and luteolin phytocon-
stituents on genotoxicity induced by halogenated boroxine [K2(B3O3F4OH)] in
human lymphocytes in vitro. Luteolin (3′,4′,5,7-tetrahydroxyflavone) is a yellow
pigmented flavone predominantly seen in the leaves of Ficus carica and to a lesser
proportion in the fruit figs chemical structure is depicted in Fig. 26.3. Chromosomal
aberrations were analyzed by the cytokinesis-block micronucleus cytome assay in
human peripheral blood lymphocytes. Luteolin inhibits genotoxic effects by
decreasing cellular proliferation and also induces cell cycle arrest in G1/S and G2/M
checkpoints (Hadžić et al., 2015).
26 Genotoxic and Antimutagenic Activity of Ficus carica Extracts 591

Fig. 26.3 Structure of luteolin found in Ficus carica leaves and fruit (National centre for biotech-
nology information (2021) Pub Chem ID:5280445)

6 Conclusion

Plant phytochemicals are the crucial source for identifying and isolating lead com-
pounds that can be formulated into a novel therapeutic agent that can be used to cure
all kinds of diseases. Evidence from ethnobotanical and pharmacological studies
reveals that the administration of natural products as medicine or food/diet supple-
ments is inversely related to disease progression. They represent a promising thera-
peutic strategy for cancer prevention. However, as they are intended to be used as
novel drugs, they must be tested for their toxicological parameters and deemed safe.
The genotoxicity, reproductive toxicology, mutagenicity testing must be performed
using a battery of in vivo and in vitro experiments authorized by a regulatory agency.
Anitmutagenicity tests commonly use Chromosome aberration assay, micronuclei
test, and comet assay. Of these, comet assay is most preferred due to ease of avail-
ability and reliability. The presence of flavonoids and polyphenols might be regarded
as the prominent phytochemicals crucial for varied therapeutic uses. With the advent
of modern technologies, several strategies can be applied for the toxicological
assessment of folklore medicine, the most powerful being the genomic approach. It
serves as a powerful tool acknowledging the pharmaco-toxicological activities of
folklore medicinal products. The involvement of “omics” technologies provides
ample opportunity and techniques to elucidate the underlying mechanisms of vari-
ous phytochemicals and create a comprehensive basis for better hazard identifica-
tion and more relevant genotoxicity assessment.

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Chapter 27
Composition and Biological Activities
of Ficus carica Latex

Mostafa M. Hegazy, Reham Hassan Mekky, Wael M. Afifi,

Ahmad E. Mostafa, and Hatem S. Abbass

Abbreviations

C NMR
13
Carbon-13 Nuclear Magnetic Resonance
1
H NMR Proton Nuclear Magnetic Resonance
ACE Angiotensin I-Converting Enzyme
AGS 6-O-acyl-β-D-glucosyl-β-sitosterol
CAE Chlorogenic Acid Equivalents
CE Catechin Equivalents.
Cv. Cultivar
EI-MS Electron Ionization Mass Spectrometry
FLP-1 Ficus Latex Peptide-1
FLP-2 Ficus Latex Peptide-2
FLP-3 Ficus Latex Peptide-3
FP1 Ficus Peroxidase 1
FP2 Ficus Peroxidase 2
FP3 Ficus Peroxidase 3
FPI Ficus Cysteine Protease I
FPII Ficus Cysteine Protease II
FPIII Ficus Serine Protease III

M. M. Hegazy (*)
Department of Pharmacognosy and Medicinal Plants, Faculty of Pharmacy,
Al-Azhar University, Cairo, Egypt
e-mail: [email protected]
R. H. Mekky
Department of Pharmacognosy, Faculty of Pharmacy, Egyptian Russian University,
Badr City, Cairo, Egypt
e-mail: [email protected]
W. M. Afifi · A. E. Mostafa · H. S. Abbass
Department of Pharmacognosy and Medicinal Plants, Faculty of Pharmacy,
Al-Azhar University, Cairo, Egypt
e-mail: [email protected]; [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 597
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_27
598 M. M. Hegazy et al.

GAE Gallic Acid Equivalents

GC-IT-MS Gas Chromatography Ion Trap Mass Spectrometry
GC-MS Gas Chromatography-Mass Spectrometry
HPLC High-Performance Liquid Chromatography
HPLC-DAD High-Performance Liquid Chromatography- Diode-Array Detector
LC-MS/MS Liquid Chromatography-Tandem Mass Spectrometry
MeOH Methanol
pI Isoelectric Point
PLCP Papain-Like Cysteine Proteases
PR-4/type II Pathogenesis-Related Protein 4 type II
SARS-CoV-2 Severe Acute Respiratory Syndrome Coronavirus 2
TFC Total Flavonoid Content
TPC Total Phenolic Content

1 Introduction

Latex is a natural sticky polymer from different plant parts and contains various
phytochemicals and antioxidants. It shows diverse biological activities such as anti-
fungal, antibacterial, antiviral, anti-inflammatory, and anticancer (Santos & Van
Ree, 2011; Upadhyay, 2011). All Ficus spp. possess latices within their vascula-
tures, affording protection and self-healing from physical assaults (Lansky et al.,
2008). Latex had been reported as a treatment for warts (Mabberley, 1997; Martínez,
2008). Although latex is not ingested per se, some precautions regarding its direct
use are needed because it is known to have keratolytic and corrosive properties
(Hemmatzadeh et al., 2003). Latex is used as expectorant, diuretic, anthelmintic and
anemia (Kirtikar & Basu, 1935). Interestingly, latex is naturally occurring in
Laticifers, defined as cells or series of connected cells containing a fluid called latex
and forming systems that permeate various tissues of the plant body (Evert, 2006).
Latex consists of a cytoplasmatic fluid of laticiferous tissues containing the usual
organelles of plant cells, such as nuclei, mitochondria, vacuoles, and ribosomes
(Kim et al., 2008). It is an aqueous suspension of a complex mixture of molecules
found in specialized secretory cells of plants, known as laticifers, which synthesize
and store a diversity of primary metabolites such as enzymatic and non-enzymatic
proteins and secondary metabolites of different classes such as flavonoids, phenolic
acids, coumarins, xanthones, terpenoids, and sterols in appreciable amounts
(Agrawal & Konno, 2009). A laticifer is a plant cell unique in shape, differentiation,
and physiological function, and its cytoplasm is a sticky fluid called latex (Kitajima
et al., 2018). In addition to the industrial importance of plant latex as a rubber
source, latex is a component of plant defense against microbes and herbivores.
Laticifers form long tubular or branched structures running throughout the plant’s
body. Owing to this structure, much latex is exuded from the cut site when the plant
body is cut, and toxic proteins and metabolites attack pests. Even though latices
share a common biological role in pest defense, their protein, and chemical
27 Composition and Biological Activities of Ficus carica Latex 599

constituents are highly variable among plant species (Hagel et al., 2008; Konno,
2011). Similarly, protein constituents of latices are variable even among organs in a
single species (Kitajima et al., 2012, 2013). Latex of F. carica (Fig. 27.1) contains
volatile components; aldehydes such as Pentanal, hexanal, heptanal, and benzalde-
hyde; alcohols such as 1-butanol-3-methyl, 1-butanol-2-methyl, and 1-heptanol;
ketones such as 6-methyl-5-hepten-2-one; monoterpenes such as α-pinene, β-pinene,
limonene, and eucalyptol; sesquiterpenes such as α-guaiene, α-bourbonene,
β-caryophyllene, and germacrene D; and miscellaneous such as methyl salicylate,
quinoline, and psoralene (Oliveira et al., 2010b). It also contains non-volatile com-
ponents; organic acids such as oxalic, citric, quinic, and shikimic (Oliveira et al.,
2010b); fatty acids such as myristic, palmitic, stearic, oleic, and linoleic (Oliveira
et al., 2010a); triterpenoids such as lupeol, lupeol acetate, and β-amyrin; steroids
such as β-sitosterol, 6-O-palmitoyl-β-D-glucosyl-β-sitosterol, and 6-O-linoleyl-β-
D-glucosyl-β-sitosterol (Oliveira et al., 2010a); and amino acids such as leucine,
tryptophan, serine, ornithine, tyrosine, and cysteine (Oliveira et al., 2010a). Ficus
carica latex contains large amounts of isoforms of ficin, a cysteine protease, which
is toxic to the caterpillars of Lepidoptera (Konno et al., 2004), and fungi
(Karnchanatat et al., 2011; López-García et al., 2012), as well as isoforms of trypsin
inhibitor, which is also known to be toxic to insects (Hilder et al., 1987), and fungi
(Huynh et al., 1992; Terras et al., 1993). Ficins to trypsin inhibitor ratio differed
between latices from immature fruit and young petioles, suggesting that, despite
these organs being young and unlignified, their latices have adopted different
defense strategies. By comparing the proteomes, metabolomes, and transcriptomes
in the latex of various F. carica organs to find out the diversity of defense strategies,
three different organs containing latex have been selected; immature fruit, which are
economically important as food; young and unlignified petioles, whose laticifers are
expected to be connected to those in leaf veins; and > 1-year-old trunks, which are
lignified and thus may have different pests from unlignified organs, Trypsin inhibi-
tors were the most abundant proteins in petiole latex, while cysteine proteases
(ficins) were the most abundant in immature fruit and trunk latices (Kitajima et al.,
2018). The expression levels of pathogenesis-related proteins were highest in the

Fig. 27.1 Ficus carica

latex
600 M. M. Hegazy et al.

latex of trunk, suggesting that this latex had adapted a defensive role against microbe
attacks (Kitajima et al., 2018). Although young petioles and immature fruit are both
unlignified soft organs, and potential food for herbivorous insects, unigenes for the
sesquiterpenoid pathway, which likely produces defense-associated volatiles, and
the phenylpropanoid pathway, which produces toxic furanocoumarins, were
expressed less in immature fruit latex (Kitajima et al., 2018).

2 Bioactive Metabolites Described in Fig (Ficus carica

L.) Latex

A wide range of bioactive metabolites was observed in F. carica L. latex, indicating

a new insight into the importance of fig latex as an undervalued source of bioactive
metabolites. In this sense, the utilization of state-of-art techniques such as gas chro-
matography (GC) and/or high-performance liquid chromatography (HPLC) cou-
pled with high-resolution mass spectrometry (MS) and tandem mass spectrometry
(MS/MS) allowed the annotation of many phenolic compounds, terpenoids, sterols,
xanthones, alkanes, alkenes, alkynes, alcohols, aldehydes, ketones, ethers, fatty
acids, aromatic compounds, amino acids, and nitrogenous compounds. Generally,
latex was collected manually from either the fruits or the leaves and stored in sterile
containers till analysis (Oliveira et al., 2010a, b; Menichini et al., 2012; Abdel-Aty
et al., 2019). It bears noting that (Menichini et al., 2012) conducted a study on the
latex collected from both the fruits and the leaves of the Italian cultivar Dottato
where the latex was collected from the leaves in June, July, and September and from
the fruits in June and July (Table 27.1 and 27.2). The process of metabolites extrac-
tion from fig latex was performed by applying multiple solvents extraction and frac-
tionation (Lazreg-Aref et al., 2012a; Menichini et al., 2012; Abdel-Aty et al., 2019;
Paşayeva et al., 2020), solid-phase extraction (Paşayeva et al., 2020) and/or, head-
space solid-phase microextraction (Oliveira et al., 2010b), Table 27.2. Totally, 130
metabolites were reported in fig latex.

2.1 Phenolic Compounds

The detected phenolic compounds in fig latex were classified into hydroxybenzoic
acids, hydroxycinnamic acids, flavonoids, and coumarins, considering each candi-
date’s name, molecular formula, occurrence, geographical origins, extraction, and
analytical methods. Table 27.2 and Fig. 27.2 illustrate the observed phenolic com-
pounds in fig latex. Besides identifying compounds, total phenolic content (TPC)
and total flavonoid content (TFC) of latex were estimated in different organs, sea-
sons, and geographical origins, as shown in Table 27.1.
27 Composition and Biological Activities of Ficus carica Latex 601

Table 27.1 Total phenolic and total flavonoid contents of different latices
Extraction
Geographical procedure after
Sample origin latex collection TPC TFC Reference
Leaves of Calabria, (Acetone: 34 ± 0.7a mg – Menichini
Cv. Dottato Italy methanol: water: CAE/ g extract et al.
(June) acetic acid, (2012)
40:40:20:0.1, v/v)
followed by
heating and
sonication
Leaves of Calabria, (Acetone: 26 ± 0.6a mg – Menichini
Cv. Dottato Italy methanol: water: CAE/g extract et al.
(July) acetic acid, (2012)
40:40:20:0.1, v/v)
followed by
heating and
sonication
Leaves of Calabria, (Acetone: 36 ± 0.8a mg – Menichini
Cv. Dottato Italy methanol: water: CAE/g extract et al.
(September) acetic acid, (2012)
40:40:20:0.1, v/v)
followed by
heating and
sonication
Fruits of Cv. Calabria, (Acetone: 29 ± 0.6a mg – Menichini
Dottato Italy methanol: water: CAE/g extract et al.
(June) acetic acid, (2012)
40:40:20:0.1, v/v)
followed by
heating and
sonication
Fruits Al Sharqia Hydroalcoholic 50.20 ± 2.50b 12.50 ± 1.10c Abdel-Aty
government, extract (80% mg GAE/g mg CE/ g et al.
Egypt MeOH) of extract extract (2019)
defatted latex
Unripe fruits Adana, Methanol extract 1.879 ± 0.004b 2.965 ± 0.006c Paşayeva
of F. carica Turkey mg GAE/ g mg CE/ g et al.
subsp. extract extract (2020)
Carica
Unripe fruits Adana, n-hexane fraction 2.910 ± 0.005b 2.227 ± 0.001c Paşayeva
F. carica Turkey of the methanol mg GAE/ g mg CE/ g et al.
subsp. extract extract extract (2020)
Carica
Unripe fruits Adana, n-hexane-ethyl 1.016 ± 0.003b 2.914 ± 0.020c Paşayeva
F. carica Turkey acetate fraction of mg GAE/g mg CE/ g et al.
subsp. the methanol extract extract (2020)
Carica extract
(continued)
602 M. M. Hegazy et al.

Table 27.1 (continued)

Extraction
Geographical procedure after
Sample origin latex collection TPC TFC Reference
Unripe fruits Adana, Methanol fraction 11.297 ± 0.001b 3.063 ± 0.011c Paşayeva
F. carica Turkey of the methanol mg GAE/g mg CE/ g et al.
subsp. extract extract extract (2020)
Carica
Unripe fruits Adana, Methanol 26.816 ± 0.007b 8.615 ± 0.009c Paşayeva
F. carica Turkey followed by mg GAE/g mg CE/ g et al.
subsp. solid-phase extract extract (2020)
Carica extraction
Leaves and Jalgaon, India Latex was diluted 98.75b μg GAE/ – Borase
fruits in water (1:5, v/v) ml et al.
(2013)
a
CAE values are chlorogenic acid equivalents
b
GAE values are gallic acid equivalents
c
(CE) values are catechin equivalents.

2.1.1 Phenolic Acids

The reported phenolic acids in fig latex were grouped into hydroxybenzoic acids
(compounds; 1–5, Table 27.2 and Fig. 27.2) and hydroxycinnamic acids (com-
pounds; 6–11, Table 27.2 and Fig. 27.2). Hydroxybenzoic acids were classified into
monohydroxybenzoic acids, dihydroxybenzoic acids, including O-methylated deriv-
atives, and trihydroxybenzoic acids conjugation with quinic acid (Oliveira et al.,
2010b; Lazreg Aref et al., 2011a; Tezcan et al., 2015; Abdel-Aty et al., 2019; Paşayeva
et al., 2020). Regarding hydroxycinnamic acids, they were either free (p-coumaric
acid, caffeic acid, ferulic acid, and sinapic acid) or conjugated with quinic acid (chlo-
rogenic acid) or 3,4-dihydroxyphenyllactic acid (Table 27.2 and Fig. 27.2).

2.1.2 Flavonoids and Coumarins

Five flavonoid derivatives were observed in fig latex (compounds 12–16, Table 27.2
and Fig. 27.2). They were analyzed either by HPLC or LC-MS/MS. They repre-
sented different flavonoids subclasses as flavones (luteolin), flavonols (quercetin
and rutin), flavan-3-ols (catechin), and anthocyanins (5-carboxypryano-pelargonidin
3-O-β-glucopyranoside). As for coumarins (compounds 17–18), psoralen and ber-
gapten were observed.

2.2 Terpenoids and Sterols

Qualitatively, terpenoids were the primary class of the reported metabolites in fig
latex (36 compounds), being classified into monoterpenes (10), sesquiterpenes (11),
and triterpenes (15), Table 27.2 and Fig. 27.2. Terpenoids possess several biological
Table 27.2 Phenolic compounds, terpenoids, and sterols reported in fig latex
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex Origin latex collection analysis Reference
Phenolic compounds: Hydroxybenzoic acids
1 Methyl salicylate C8H8O3 Fruits Al Sharqia Hydroalcoholic extract (80% GC-MS Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
fruits from F. carica region, Portugal Microextraction (HS-SPME). (2010b)
Cv. Pingo de Mel
2 Protocatechuic acid C7H6O4 Unripe fruits Sousse, Tunisia n-hexane and n-hexane -ethyl LC-MS/MS Lazreg Aref
acetate fractions of the et al. (2011a)
methanolic extract
Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
Leaves Bursa, Turkey Water fraction after defatting HPLC Tezcan et al.
and multiple solvents (2015)
fractionation
3 Vanillic acid C8H8O4 Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
27 Composition and Biological Activities of Ficus carica Latex

4 Methyl vanillate C9H10O4 government, MeOH) of defatted latex et al. (2019)

Egypt
5 Trigalloylquinic acid C28H24O18 Unripe fruits F. Adana, Turkey Methanol followed by LC-MS/MS Paşayeva
carica subsp. carica solid-phase extraction et al. (2020)
Phenolic compounds: Hydroxycinnamic acids
(continued)
603
Table 27.2 (continued)
604

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex Origin latex collection analysis Reference
6 p-coumaric acid C9H8O3 Unripe fruits Sousse, Tunisia n-hexane fraction of the LC-MS/MS Lazreg Aref
methanolic extract et al. (2011a)
Leaves Bursa, Turkey Water fraction after defatting HPLC Tezcan et al.
and multiple solvents (2015)
fractionation
7 Caffeic acid C9H8O4 Unripe fruits Sousse, Tunisia n-hexane and n-hexane -ethyl LC-MS/MS Lazreg Aref
acetate fractions of the et al. (2011a)
methanolic extract
Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt.
Leaves Bursa, Turkey Water fraction after defatting HPLC Tezcan et al.
and multiple solvents (2015)
fractionation
8 Chlorogenic acid C16H18O9 Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
9 Ferulic acid C10H10O4 Unripe fruits Sousse, Tunisia n-hexane and n-hexane-ethyl LC-MS/MS Lazreg Aref
acetate fractions of the et al. (2011a)
methanolic extract
Fruits Al Sharqia hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
Leaves Bursa, Turkey Water fraction after defatting HPLC Tezcan et al.
and multiple solvents (2015)
fractionation
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex Origin latex collection analysis Reference
10 Sinapic acid C11H12O5 Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
11 Rosmarinic acid C18H16O8 Unripe fruits F. Adana, Turkey Methanol followed by LC-MS/MS Paşayeva
carica subsp. carica solid-phase extraction et al. (2020)
Phenolic compounds: Flavonoids
12 Luteolin C15H10O6 Unripe fruits Sousse, Tunisia n-hexane and n-hexane -ethyl LC-MS/MS Lazreg Aref
acetate fractions of the et al. (2011a)
methanolic extract
13 Quercetin C15H10O7 Leaves Bursa, Turkey Water fraction after defatting HPLC Tezcan et al.
and multiple solvents (2015)
fractionation
14 Quercetin 3-O-rutinoside (rutin) C27H30O16 Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
15 Catechin C15H14O6 Fruits Al Sharqia Hydroalcoholic extract (80% HPLC Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
16 5-carboxypryano-pelargonidin C24H21O12+ Unripe fruits F. Adana, Turkey Methanol followed by LC-MS/MS Paşayeva
3-O-β-glucopyranoside carica subsp. carica solid-phase extraction et al. (2020)
27 Composition and Biological Activities of Ficus carica Latex

Phenolic compounds: Coumarins

(continued)
605
Table 27.2 (continued)
606

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex Origin latex collection analysis Reference
17 Psoralen C11H6O3 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Fruits Al Sharqia Hydroalcoholic extract (80% GC-MS Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
fruits from F. carica region, Portugal Microextraction (HS-SPME) (2010b)
Cv. Pingo deMel
18 Bergapten C12H8O4 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Terpenoids: Monoterpenes
19 α-thujene C10H16 Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
20 α-pinene C10H16 fruits from F. carica region, Portugal Microextraction (HS-SPME) (2010b)
21 β-pinene C10H16 Cv. Pingo de Mel
22 Limonene C10H16
23 Terpinolene C10H16
24 3-Bornanone C10H16O Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex Origin latex collection analysis Reference
25 Eucalyptol C10H18O Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
26 Linalool C10H18O fruits from F. carica region, Portugal Microextraction (HS-SPME) (2010b)
27 cis-linalool oxide C10H18O2 Cv. Pingo de Mel
28 Epoxylinalol C10H18O2
Terpenoids: Sesquiterpenes
29 Epi-bicyclosesquiphellandrene C15H24 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
30 Germacrene-D C15H24 Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
fruits from F. carica region, Portugal Microextraction (HS-SPME). (2010b)
Cv. Pingo de Mel
31 Delta-Cadinene C15H24 Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
fruits from F. carica region, Portugal Microextraction (HS-SPME) (2010b)
Cv. Pingo de Mel
27 Composition and Biological Activities of Ficus carica Latex

32 α-guaiene C15H24 Immature green Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
33 α-bourbonene C15H24 fruits from F. carica region, Portugal Microextraction (HS-SPME) (2010b)
34 α -caryophyllene C15H24 Cv. Pingo de Mel
35 β-caryophyllene C15H24
36 τ-muurolene C15H24
37 α-calacorene C15H24
38 Trans-α-bergamotene C15H24
(continued)
607
Table 27.2 (continued)
608

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex Origin latex collection analysis Reference
39 Aristolone C15H22O Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
Terpenoids: Triterpenes
40 Squalene C30H50 Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
41 1, 6, 10, 14, 18, C30H50O inedible unripe fruit Tunisia et al. (2012a)
22-Tetracosahexaen-3-ol
42 β-Amyrin C30H50O Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
43 α-Amyrin C30H50O Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
44 α-Amyrenyl acetate C32H52O2 Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
45 Maragenin I acetate C31H48O3 inedible unripe fruit Tunisia et al. (2012a)
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex Origin latex collection analysis Reference
46 Lupeol (Lup-20(29)-en-3-ol C30H50O Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(3β)-) (June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal
n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
1
47 Lupeol acetate C32H52O2 Green fruits (July) Sari, Mazandaran n-hexane followed by H, 13C NMR, Jeivad et al.
Province, Iran chloroform EI-MS (2020)
Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
1
48 Lupeol palmitate C46H80O2 Green fruits in July Sari, Mazandaran n-hexane followed by H, 13C NMR, Jeivad et al.
2015 Province, Iran chloroform EI-MS (2020)
49 D:C-Friedours-7-en-3-ol, (3β)- C30H52O Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
27 Composition and Biological Activities of Ficus carica Latex

(June, July,
September)
50 Olean-12-en-3-ol, acetate, (3β)- C32H52O2 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
(continued)
609
Table 27.2 (continued)
610

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex Origin latex collection analysis Reference
51 Urs-12-en-24-oic acid, 3-oxo-, C31H48O3 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
methyl ester, (+)- (June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
52 A-Neooleana-3(5), 12-diene C30H48 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
53 A′-Neogammacer-22(29)-en- C32H52O2 Fruits of Cv. Dottato
3-ol, acetate, (3β, 21β)- (June, July)
Leaves of Cv. Dottato
(June, July,
September)
54 Betulol C30H50O2 Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
Sterols
55 Lanosterol C30H50O Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex Origin latex collection analysis Reference
56 Lanosta-8,24-dien-3-ol,acetate(3β)- C32H52O2 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
inedible unripe fruit Tunisia et al. (2012a)
57 5α-Lanosta-7,24-dien-3β-ol C32H52O2 Fruits of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
acetate (June, July) n-hexane et al. (2012)
Leaves of Cv. Dottato
(June, July,
September)
58 5α-Lanosta-9(11), 25-dien-3β-ol C34H56O2 Fruits of Cv. Dottato
acetate (June, July)
Leaves of Cv. Dottato
(June, July,
September)
59 β-Sitosterol C29H50O Fruits of Cv. Dottato
(June, July)
Leaves of Cv. Dottato
27 Composition and Biological Activities of Ficus carica Latex

(June, July,
September)
Green fruits Mirandela Different extracts of GC-IT-MS/ Oliveira et al.
region, Portugal n-hexane, dichloromethane, HPLC-DAD (2010a)
alkaline hydrolysis
60 6-aza-B-Homo-5αcholestane C27H49N Caprifig Jrani Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
61 9, 19-Cyclolanost-24-en-3-ol C30H50O inedible unripe fruit Tunisia et al. (2012a)
611
612 M. M. Hegazy et al.

(1) R1=R2=H (2) (5) R1=R2=R3=H (6) R=H (12)

R1=CH3 R2=H (3) R1=OH R2=R3=H (7) R=OH (13)
R1=R2=CH3 (4) R1=OH R2=H R3=quinic acid (8) R=O-rutinoside (14)
R1=OCH3 R2=R3=H (9)
R1=R2=OCH3 R3=H (10)
R1=R2=H
R3=3,4-dihydroxyphenyl-acetic acid
(11)

(15) (16) (17) (18)

(19) (20) (21) (22) (23) (24) (25) (26) (27) (28) (29) (30)

(31) (32) (33) (34) (35) (36) (37) (38)

(39) (40) (41) (42)

R=H (43) R= (45) R=H (46) (49) (50)

R=acetate (44) R=acetate (47)
R=palmitate (48)

Fig. 27.2 Chemical structures of phenolic compounds, terpenoids, and sterols r eported in the
latex of figs
27 Composition and Biological Activities of Ficus carica Latex 613

(51) (52) (53) (54)

R=H (55) (57) (58) (59)

R= acetate (56)

(60) (61)

Fig. 27.2 (continued)

activities, and hence, their presence in the latex of figs contributes significantly to
enhancing its biological potential (Wang et al., 2005). Besides, sterols occurrence
was observed with seven derivatives (Table 27.2 and Fig. 27.2).

2.3 Xanthones, Hydrocarbons, Fatty Acids, Aromatic

Compounds, Amino Acids, and Other
Nitrogenous Compounds

Xanthones were represented by two derivatives, namely xanthotoxin and mangif-

erin which could be an addition to the biological potential of fig latex. Xanthones
have a wide range of biological activities such as anti-inflammatory, anti-cancer,
anti-Alzheimer, and cardioprotective (El-Seedi et al., 2010). Also, several hydrocar-
bons were reported. They are classified into alkanes (8), alkenes (2), alkynes (1),
alcohols (7), aldehydes (5), ketones (3), and ethers (1). Aromatic compounds were
present with eight derivatives. Besides, fatty acids were widely present with 16
derivatives, including unsaturated fatty acids incorporating the antihyperlipidemic
and cardioprotective activity of fig latex (Oliveira et al., 2010a). In line with nitrog-
enous compounds, 16 derivatives were reported in fig latex with essential amino
acids, providing a new nutritive prospect for such an undervalued plant part (Oliveira
et al., 2010a), Table 27.3 and Fig. 27.3.
Table 27.3 Xanthones, alkanes, alkenes, alkynes, alcohols, aldehydes, ketones, ethers, fatty acids, aromatic compounds, amino acids, and nitrogenous
614

compounds reported in fig latex

Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex origin latex collection analysis Reference
Xanthones
62 Xanthotoxin C16H8O4 Fruits Al Sharqia Hydroalcoholic extract (80% GC-MS Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
63 Mangiferin C9H18O11 Unripe fruits F. carica Adana, Turkey n-hexane-ethyl acetate LC-MS/MS Paşayeva
subsp. carica fraction of the methanol et al. (2020)
extract
Alkanes
64 Pentadecane C15H32 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
65 Eicosane C20H42 Leaves of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June) n-hexane et al. (2012)
66 Tricosane C23H48 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
67 Hexacosane C26H54 unripe fruit Tunisia et al. (2012a)
68 Heptacosane C27H56 Leaves of Cv. Dottato Calabria, Italy Methanol followed by GC-MS Menichini
(June) n-hexane et al. (2012)
69 Octacosane C28H58 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
70 Nanocosane C29H60
71 Triacontane C30H62
Alkenes
72 2, 6, 10-Dodecatrien-1-ol C12H20O Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
73 1, E-8, Z-10-Hexadecatriene C16H28 unripe fruit Tunisia et al. (2012a)
Alkynes
74 7-Pentadecyne C15H28 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex origin latex collection analysis Reference
Alcohols
75 1-butanol-2-methyl C5H12O Immature green fruits Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
76 1-butanol-3-methyl C5H12O from F. carica cultivar region, Portugal Microextraction (HS-SPME) (2010b)
77 1-pentanol C5H12O Pingo de Mel
78 1-hexanol C6H14O
79 1-heptanol C7H16O
80 Phenylethyl alcohol C8H10O
81 Phenylpropyl alcohol C9H12O
Aldehydes
82 Pentanal C5H10O Immature green fruits Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
83 Hexanal C6H12O from F. carica cultivar region, Portugal Microextraction (HS-SPME) (2010b)
84 Heptanal C7H14O Pingo de Mel
85 Octanal C8H16O
86 Benzaldehyde C7H6O
Ketones
87 6-methyl-5-hepten-2-one C8H14O Immature green fruits Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
from F. carica cultivar region, Portugal Microextraction (HS-SPME) (2010b)
Pingo de Mel
88 Dodecan-2-on C11H18O2 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
27 Composition and Biological Activities of Ficus carica Latex

unripe fruit Tunisia et al. (2012a)

89 2,5,8-Trimethyl-1-tetralone C13H16O Fruits Al Sharqia Hydroalcoholic extract (80% GC-MS Abdel-Aty
government, MeOH) of defatted latex et al. (2019)
Egypt
Ethers
90 Oxirane C2H4O Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
Fatty acids
(continued)
615
Table 27.3 (continued)
616

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex origin latex collection analysis Reference
91 Myristic acid C14H28O2 Green fruits Mirandela Methanol followed by GC-IT-MS Oliveira et al.
92 Pentadecanoic acid C15H30O2 region, Portugal alkaline hydrolysis and (2010a)
derivatization
93 Palmitic acid (Hexadecanoic C16H32O2 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
acid) unripe fruit Tunisia et al. (2012a)
94 Heptadecanoic acid C17H34O2 Green fruits Mirandela Methanol followed by GC-IT-MS Oliveira et al.
region, Portugal alkaline hydrolysis and (2010a)
95 cis-10-heptadecenoic acid C17H32O2
derivatization
96 Stearic acid C18H36O2
97 Oleic acid C18H34O2
98 Elaidic acid C18H34O2
99 Linoelaidic acid C18H32O2 Unripe fruits F. carica Adana, Turkey Methanol followed by LC-MS/MS Paşayeva
subsp. carica solid-phase extraction et al. (2020)
100 Linoleic acid C18H32O2 Green fruits Mirandela Methanol followed by GC-IT-MS Oliveira et al.
region, Portugal alkaline hydrolysis and (2010a)
derivatization
101 Cis-Linoleic acid methyl ester C19H34O2 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
102 Arachidic acid C20H40O2 Green fruits Mirandela Methanol followed by GC-IT-MS Oliveira et al.
103 Heneicosanoic acid C21H42O2 region, Portugal alkaline hydrolysis and (2010a)
104 Behenic acid C22H44O2 derivatization
105 Tricosanoic acid C23H46O2
106 Lignoceric acid C24H48O2
Aromatic compounds
107 1,8-Dimethoxynaphthalene C12H12O2 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
unripe fruit Tunisia et al. (2012a)
M. M. Hegazy et al.
Molecular Occurrence of the Geographical Extraction procedure after Method of
# Compound formula latex origin latex collection analysis Reference
108 6-methyl 4-chromanone C10H10O2 Unripe fruits F. carica Adana, Turkey Methanol followed by LC-MS/MS Paşayeva
subsp. carica solid-phase extraction et al. (2020)
n-hexane-ethyl acetate
fraction of the methanol
extract
109 p-OH-Phenylacetic acid C8H8O2 Unripe fruits Sousse, Tunisia n-hexane fraction of the LC-MS/MS Lazreg Aref
methanolic extract et al. (2011a)
110 Oxalic acid, C14H18O4 Unripe fruits F. carica Adana, Turkey n-hexane-ethyl acetate LC-MS/MS Paşayeva
2-methylphenylpentyl ester subsp. carica fraction of the methanol et al. (2020)
extract
111 Phthalic acid C8H6O4 Fruits Al Sharqia Hydroalcoholic extract (80% GC-MS Abdel-Aty
112 Dibutyl phthalate C16H22O4 government, MeOH) of defatted latex et al. (2019)
Egypt.
113 bis (2-Ethylhexyl) phthalate C24H38O4 Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
114 [3.2] Metacyclophane-10-ene C16H14 unripe fruit Tunisia et al. (2012a)
Amino acids
115 Argenine C6H14N4O2 Unripe fruits Sousse, Tunisia n-hexane and n-hexane-ethyl LC-MS/MS Lazreg Aref
acetate fractions of the et al. (2011a)
methanolic extract
27 Composition and Biological Activities of Ficus carica Latex

(continued)
617
Table 27.3 (continued)
618

Molecular Occurrence of the Geographical Extraction procedure after Method of

# Compound formula latex origin latex collection analysis Reference
116 Glycine C2H5NO2 Green fruits Mirandela 0.1M HCl HPLC-DAD Oliveira et al.
117 Alanine C3H7NO2 region, Portugal (2010a)
118 Serine C3H7NO2
119 Asparagine C4H8N2O3
120 Ornithine C5H12N2O2
121 Glutamine C5H10N2O3
122 Leucine C6H13NO2
123 Tryptophan C11H12N2O2
124 Phenylalanine C9H11NO2
125 Tyrosine C6H14N2O2
126 Histidine C6H9N3O2
127 Lysine C9H11NO3
128 Cysteine C3H7NO2S
Nitrogenous compounds
129 4-(4-Methoxyphenyl)-2,6- C24H19NO Caprifig Jrani inedible Mesjed Aissa, n-hexane GC-MS Lazreg-Aref
diphenylpyridine unripe fruit Tunisia et al. (2012a)
130 Quinoline C9H7N Immature green fruits Mirandela Headspace Solid-Phase GC-IT-MS Oliveira et al.
from F. carica cultivar region, Portugal Microextraction (HS-SPME). (2010b)
Pingo deMel
M. M. Hegazy et al.
27 Composition and Biological Activities of Ficus carica Latex 619

(62) (63) (64) (65)

(66) (67) (68) (69)

(70) (71) (72) (73)

(74) (75) (76) (77) (78) (79)

(80) (81) (82) (83) (84) (85) (86)

(87) (88) (89) (90)

(91) (92) (93) (94)

(95) (96) (97) (98)

(99) R=H (100) (102) (103)

R= CH3 (101)

(104) (105) (106) (107)

(108) (109) (110) R=H (111)

R=butyl (112)
R= 2-ethylhexyl (113)

Fig. 27.3 Chemical structures of xanthones, hydrocarbons, fatty acids, aromatic compounds,
amino acids, and other nitrogenous compounds
(continued)
620 M. M. Hegazy et al.

(114) (115) (116) (117)

(118) (119) (120) (121)

(122) (123) R=H (124) (126)

R=OH (125)

(127) (128) (129) (130)

Fig. 27.3 (continued)

2.4 Rubber

Natural rubber is a type of crucial biopolymer with meaningful use and strategic
importance in human society, composed of 320–35,000 isoprene molecules (cis-1,4-
polyisoprene) (Kang et al., 2000); of which the constituent monomer isopentenyl
pyrophosphate (IPP) is synthesized from the mevalonate (MVA) pathways (Men
et al., 2018). Rubber is produced in latex at the expense of high-energy cost and is
considered a secondary metabolite with no known function in plant cells. Although
it is not fully understood why plants produce rubber, it has been suggested that the
latex secretion is a defense against wounding and/or predators such as insects and
microorganisms (Kim et al., 2003) and is widely used in the manufacture of various
industrial and medicinal products (Kang et al., 2000), have unique properties which
include resilience, elasticity, abrasion and impact resistance, efficient heat disper-
sion, and malleability at cold temperatures. As a result, natural rubber is irreplace-
able in many applications (heavy-duty tires, medical devices, surgical gloves, etc.)
(Men et al., 2018). Therefore, to understand the molecular mechanism of rubber
biosynthesis, it is helpful to investigate other rubber-producing plants and character-
ize and compare the enzymes related to rubber biosynthesis in the different plant
species. Recently, we have identified the fig tree (Ficus carica) as an alternative
rubber-producing temperate plant and reported that the high level of divalent metal
ion in the latex serum of the fig tree exerts different rubber biosynthetic activities in
F. carica. The amounts of latex in plants and latex rubber content vary depending on
the physiological conditions of plants influenced by environments. The molecular
size of rubber is determined by the action of enzymes and/or factors such as rubber
transferase and rubber elongation factor (Kang et al., 2000).
27 Composition and Biological Activities of Ficus carica Latex 621

2.5 Proteins

Latex of F. carica is rich in proteins of different classes, biological functions, and

industrial applications (Mohammad & Alzweiri, 2021). Fig tree latex contains
between 10% and 17% enzyme and non-enzyme proteins (Lazreg-Aref et al., 2018).
Zhai et al. (2021) reported the presence of 38.9% papain-like cysteine proteases in
the total protein content as the most abundant protein of ripe fig fruit latex (Zhai
et al., 2021). Furthermore, many enzyme activities of the latex were characterized,
such as protease, peroxidase, chitinase, amylase, and lipase, which play vital roles
for the fig plant. For instance, chitinases and peroxidases serve as defensive proteins
protecting the fig plant against insects and fungi (Mohammad & Alzweiri, 2021). In
addition, many non-enzymatic peptides and proteins are involved as a part nonspe-
cific system of protection against microbes, such as a low molecular weight chitin-
binding protein of antifungal activity isolated from latex collected from young
branches of fig tree cultivated in Uzbekistan at the beginning of fruiting stage. The
antifungal protein was classified as a pathogenesis-related protein 4 type II (PR-4/
type II) according to its partial amino acid sequence and chitin binding ability
(Table 27.4) (Mavlonov et al., 2008). Moreover, three peptides of angiotensin
I-converting enzyme (ACE) inhibitory activity were isolated from stalks latex of a
fig tree grown in Japan. Its amino acid sequences were identified as Ala-Val-Asn-
Pro-Ile-Arg (FLP-1), Leu-Tyr-Pro-Val-Lys (FLP-2), and Leu-Val-Arg (FLP-3)
(Table 27.4) (Maruyama et al., 1989).

2.5.1 Protease Activity

Proteases are the part of latex responsible for breaking down proteins into smaller
polypeptides or amino acids. They are classified according to the catalytic amino
acid of the active site into four classes: cysteine proteases, serine proteases, aspartic
proteases, and metalloproteases (Mohammad & Alzweiri, 2021). Generally, plant
proteases participate in many vital functions, such as nutrient mobilization, fruit
maturation, protection of fruit during ripening against plant pathogens, and wound
healing by latex coagulation injuries caused by a biotic or abiotic agent (Mohammad
& Alzweiri, 2021; Azarkan et al., 2011). The latex of F. carica has several proteases
of two classes: cysteine and serine proteases. Most latex proteases are members of
the cysteine protease class, commonly named ficins. While collagenolytic, fibrino-
lytic, and subtilase activities are serine protease classes (Mohammad & Alzweiri,
2021). Changes in the proteolytic enzyme content all over the year reported by
Robbins, 1935 for the latex of the fig tree growing in Alabama, USA. The thin and
watery latex collected in June contained the lowest concentration of proteolytic
enzymes, then the concentration dramatically increased ten folds after 2 months
in August.
On the contrary, the thick and creamy latex collected from October to April con-
tained the highest levels of proteolytic enzymes (Robbins, 1935). The proteolytic
622 M. M. Hegazy et al.

activity of the fig latex is a summation of cysteine and serine proteases activities.
Vatić et al. (2020) showed that the fig latex has higher efficiency than papain and
trypsin in releasing Listeria monocytogenes after dissociating the fermented meat
products for enumeration of foodborne pathogenic bacteria. Furthermore, latex
activity was not affected by Triton X-100 as a strong non-ionic detergent with more
cost-efficiency than other alternatives since it could be used efficiently in concentra-
tion100 times than previously reported (Vatić et al., 2020).

Cysteine Protease Activity

Ficin or ficain (EC 3.4.22.3) is the cysteine protease of F. carica latex and is widely
present in Ficus species in many isoforms (Zhai et al., 2021). The isolated ficin
isoforms showed stability over a wide pH range with maximum catalytic activity at
pH = 7. Ficin has more than one enzyme active; it showed a protease activity and an
intrinsic peroxidase activity performed on two separate active sites (Mohammad &
Alzweiri, 2021). Ficin is a member of the papain-like cysteine proteases (PLCP)
family, characterized by strong proteolytic activity and broad substrate specificity
(Zhai et al., 2021). Ficins as a major PLCP of ripe fig fruit, present in receptacles
more than in inflorescences (Zhai et al., 2021). PLCP, including ficin, is of high
commercial value and is widely used in cheese making, meat tenderization, biscuit
baking, and producing bioactive peptides as versatile biocatalysts and digestive
drugs (Di Pierro et al., 2014; Zhai et al., 2021). Zhai et al. (2021) identified seventy-
four proteins in the latex of the fig fruit grown in China with molecular wights rang-
ing from 8.9 to 206.1 kDa and isoelectric point (pI) values of 4.84–10.7. The latex
contained trypsin-like protease inhibitor, pathogenesis-related (PR) protein iso-
forms, chitinase, and endochitinase, collectively stress-response proteins that play
an essential role in fruit protection against different plant pathogens (Zhai et al.,
2021). The unprotected cysteine proteases undergo degradation through irreversible
oxidation and/or autolysis, making chromatographic profiles complex and challeng-
ing to resolve. Azarkan et al. (2011) used a thiol-pegylation strategy by specific and
reversible monomethoxypolyethylene glycol (mPEG) reagent to purify the fully
active ficins more easily (Azarkan et al., 2011). Protection of ficins thiol functions
is a prerequisite before any fractionation to prevent autolysis and/or irreversible
oxidation of these proteases and improve chromatographic separation. Catalytic
cysteine residues of ficins are protected with Selective and reversible reagent such
as methylmethanethiol sulfonate (MMTS) (Table 27.4) (Azarkan et al., 2011).
Unlike papain, ficin is more acidic with higher substrate-binding efficiency. It was
necessary to find other ways to produce this industrially valuable protease in a way
that preserves fig tree from branch cutting. So the large-scale production of ficin
could be possible using cloned Escherichia coli BL21, which can express 0.67 mg/
mL of recombinant ficin under optimized conditions (Sattari et al., 2020).
27 Composition and Biological Activities of Ficus carica Latex 623

Serine Protease Activity

Collagenase is a serine protease that catalyzes collagen and gelatin breakdown

(Mohammad & Alzweiri, 2021). Collagenolytic activity in the latex of F. carica var.
Brown Turkey grown in Montenegro was studied by Raskovic and Polovic in
2016 in an attempt to explain its uses for skin in the Western Balkans. Collagenolytic
serine protease of the fig latex can cleave native collagen triple helix. Therefore, the
treatment with fig latex collagenase could promote connective tissue remodelings
such as debridement of wounds and ulcers. Also, collagenase improved the penetra-
tion of low molecular weight molecules across the gelatin hydrogel.
Consequently, the collagenase of fig latex could be used to enhance the delivery
of the anti-proliferative drugs in cancer treatment. Moreover, latex collected in
spring has two-fold the collagenolytic activity of the summer. In addition, the col-
lagenase of the fig latex was stable for up to 1 h in the simulated gastric conditions
(Raskovic et al., 2016). A novel collagenolytic activity was purified and character-
ized by Raskovic et al. (2014) in the latex collected from shoots of F. carica var.
brown Turkey grown in Montenegro (Table 27.4). The purified collagenase was
acidic and showed optimum catalyzing activity at pH 8.0–8.5 and 60 °C with a pI
value of about 5. Based on the specificity and high stability of the purified collage-
nase, the authors suggest that the enzyme could be useful for applications in many
fields such as biotechnology and medicine (Raskovic et al., 2014). Hamed et al.
(2020) purify and characterize a fibrinolytic serine protease (FPIII) from fruit latex
of the Egyptian F. carica (Table 27.4). Authors suggest that FPIII is responsible for
or participates in the fibrinogenolytic and anticoagulant activities that control ficin
by counteracting its procoagulant activity. It showed thermal stability up to 60 °C
and good catalytic efficiency on different protein substrates with optimum pH of 8.5
(Hamed et al., 2020).

2.5.2 Peroxidase Activity

Peroxidases are heme-containing glycoproteins that catalyze oxidation reactions.

Therefore, they are involved in plant protection against microorganisms (Mohammad
& Alzweiri, 2021). Latex collected from F. carica branches was used to purify three
peroxidase isoenzymes named FP1, FP2, and FP3. The isoenzyme FP1 was com-
pletely purified with optimum activity at pH 5.5 and 40 °C (Table 27.4). On the
other hand, FP2 and FP3 isoenzymes were partially purified because of their low
concentration and level of activity. The high concentration of calcium ions increases
the activity of FP1 and FP2 isoenzymes. The three isoenzymes showed a broad
specificity to some phenolic substrates and O-Phenylenediamine. These isoen-
zymes’ properties satisfy the prerequisites needed for industrial and environmental
applications (Elsayed et al., 2018).
624 M. M. Hegazy et al.

Table 27.4 Enzyme activities from fig latex

Name and/or
class Isolation MW References
Chitin binding Latex was processed and subjected to successive 6481 kDa Mavlonov
protein (PR-4/ chromatographic purification over chitin as an et al. (2008)
type II) affinity matrix, SP Sephadex C-50 as cation
exchanger, and reversed-phase high-performance
liquid chromatography (RP-HPLC)
FLP-1, FLP-2, The boiled latex was filtered, ultra-filtered, and 668, 618, and Maruyama
and FLP-3 fractionated over SP-Sephadex C-25 as a cation 386 Da, et al. (1989)
(ACE exchanger, and then the active fractions were respectively
inhibitory chromatographed over Sephadex LH-20
peptides) followed by RP-HPLC
Ficin A, B, C, Fresh latex was collected in the presence of About Azarkan
D1, and D2 MMTS followed by fast protein liquid 24 kDa for all et al. (2011)
(PLCP) chromatography over an SP-Sepharose as a ficin forms
cation exchanger, and then the eluted ficins were
pegylated by mPEG reagent
Collagenase Latex was centrifuged to remove insoluble gum. 41 kDa Raskovic
(serine Then, the clear water fraction was et al. (2014)
protease) chromatographed over a Sephadex G-50 column
as an ion exchanger matrix, followed by further
purification over a thiol-Sepharose column for
the fractions with caseinolytic and gelatinolytic
and the unbound fractions with collagenolytic
and gelatinolytic activities were collected
FPIII Latex fractionated over Diethylaminoethyl 48 kDa Hamed
(fibrinolytic cellulose as ion exchange resin, and the fractions et al. (2020)
serine protease) with highest proteolytic activity purified over
carboxymethyl-cellulose column followed further
purification over Sephacryl S-200 column for the
concentrated fractions with highest proteolytic
activity
FP1 isoenzyme Insoluble materials of the processed latex were 30 kDa Elsayed
(peroxidase) removed, and aqueous latex with peroxidase et al. (2018)
activity was fractionated over Carboxymethyl
(CM)-Sepharose column as a cation exchanger;
then, the fractions with peroxidase activity were
subjected to diethylaminoethyl (DEAE)-
Sepharose column as an anion-exchanger and
Sephacryl S-200 column as a size-exclusion
matrix
Lipase Latex was ultra-centrifuged and then fractionated 29 kDa Lazreg-Aref
over silica gel column chromatography eluting et al.
with acetone. Then the polarity was increased (2012b)
with water
27 Composition and Biological Activities of Ficus carica Latex 625

2.5.3 Chitinase Activity

Chitinase is the enzyme that catalyzes the breakdown of chitin, which is a structural
component of insects’ exoskeleton and fungal cell walls. Chitinase has the ability of
retarding insect development and destruction of the fungal cell wall. Therefore, chi-
tinases serve as one of the plant defensive strategies, which also could be applied as
biopesticides and antifungal agents. However, chitinases of F. carica latex were
isolated mainly from tree trunks (Mohammad & Alzweiri, 2021). The chitinolytic
activity of the latex collected from F. carica var. brown Turkey from the beginning
of the flowering to the fruit ripening stages was examined, which showed 6.5 times
higher activity at the beginning of flowering than ripened fruit. Accordingly, the
optimal fig protection against plant pathogens at the beginning of the flowering
stage depends mainly on chitinolytic activity, which is a critical period for the fig
plant and then gradually replaced over time with other strategies (Raskovic
et al., 2016).

2.5.4 Amylase Activity

Amylases are among industrially essential enzymes involved in the hydrolysis of

starch (Gupta et al., 2003). Latices collected from unripe fruit stalks of F. carica var.
Kahli and Bidhi grown in Tunisia were investigated for their amylolytic activities.
Bidhi and Kahli amylases showed optimum activity in pH of 6.5 and 7 at 45 °C and
a half-life of 85 and 60 min at 80 °C, respectively. They were highly stable in a wide
pH range (4–12). amylase activity of Bidhi variety increased 260% by addition of
ferrous ion or cupric ion and was strongly decreased by magnesium ion and
EDTA. In the same way, the presence of calcium and magnesium ions caused a
dramatic increase of Kahli amylase activity up to 220% and 260%, respectively.
Amylases of both varieties were insensitive to inhibition by glucose with high ther-
mostability in the presence of starch. Therefore, the properties of these amylases
could be very efficient for glucose production from starch (Lazreg Aref et al., 2011c).

2.5.5 Lipase Activity

Lipases are enzymes that hydrolyze oils and fats into di- and monoacylglycerol,
fatty acids, and glycerol. Lipase enzyme (ZL) was purified and characterized in
Latex collected from unripe fruit stalks of F. carica var. Zidi is grown in Tunisia
(Table 27.4). Lipase activity was 12-fold higher for long-chain triacylglycerols than
the short-chain. The purified lipase was stable in pH range 4–8 and displayed a half-
life of 90 min at 60 °C. The purified lipase activity was strongly reduced by ferrous,
magnesium, and zinc ions but significantly increased by calcium and cupric ions.
Features of purified lipase activity such as stability over a wide pH range and at high
temperature, surfactant resistance, and storage stability make it a suitable potential
626 M. M. Hegazy et al.

additive for many fields such as pharmaceutical, leather, and detergent industries
(Lazreg-Aref et al., 2012b).

3 Biological Activities of Fig Latex

The biological activities of the latex of F. carica belong to its diversity of secondary
metabolites of different chemical classes, as mentioned earlier as phenolics, flavo-
noids, terpenoids, etc. These different metabolites showed many biological activi-
ties, such as antibacterial, antifungal, antivirals, anticancer, anti-angiogenic,
antioxidant, anti-inflammatory, anticoagulant, antihypertensive, and hepatoprotec-
tive antiparasitic activities.

3.1 Antimicrobial Activity

Microorganisms are found everywhere around us in soil, water, air, skin, and most
man-made environments throughout the world. The symptoms of such infections
are generalized inflammation and sepsis, and when they occur in critical body
organs, such as the lungs, the urinary tract, and kidneys, the results can be fatal
(Al-Sokari & El Sheikha, 2015). Latex of F. carica exhibited strong activities
against tested microorganisms, including gram-positive bacteria, gram-negative
bacteria, and yeast. These activities are represented in Tables 27.5 and 27.6.
These results revealed that caprifig latex had inhibition of the growth of all bacte-
rial and yeast species represented in this chapter. The pharmacological properties
are probably due to the high content of enzymes, flavonoids, and furanocoumarins
from fig latex (Rashid et al., 2014). Antimicrobial activities of the hexane extract are
generally due to the high content of coumarins. Free 6-OH in the coumarin nucleus
is essential for antifungal activity, and the free hydroxyl group at position 7 is essen-
tial for antibacterial activity (Lazreg-Aref et al., 2012a). Antimicrobial activities are
also assigned to low molecular weight short polypeptides in fig latex, which are
shown to possess antibacterial and antifungal activities. It seems that ficin, chitin-
ases, and chitinase-like proteins abundantly accumulated in fig’s latex are the major
defense mechanism against insects and fungi rely on its proteolytic and chitinolytic,
explaining the potential antifungal activities. The antifungal activity of the isolated
protein can be explained by its chitin-binding ability as a protein binding to the
chitin component of the cell wall that suppresses fungal hyphae growth. (Mavlonov
et al., 2008; Raskovic et al., 2016). Our data showed no uniform response between
bacterial and fungal strains in terms of susceptibility to antimicrobial compounds in
different extracts; thus, different solvents have been found to have the varied capac-
ity to extract different phytoconstituents depending on their solubility or polarity in
the solvent. The ethyl acetate and chloroform fractions have a medium polarity, so
the substances that have the most important biological activities were the medium
27 Composition and Biological Activities of Ficus carica Latex 627

Table 27.5 Antibacterial activities of F. carica latex different fractions

Microorganism and Activities of
type different extracts Methodology Reference
Aeromonas Weak (L.) Well diffusion method Ahmed (2016)
hydrophila (Inhibition zone)
(Gram-negative
bacteria)
Citobacter freundei Weak (LH.) Disk diffusion assay Lazreg Aref et al. (2010,
(Gram-positive Strong (L., LC., (Inhibition zone) 2011b)
bacteria) LE.) Paper diffusion assay
(Inhibition zone)
Microwell dilution
assay (MIC)
Enterococcus fecalis Weak (LC.) Paper diffusion assay Lazreg Aref et al. (2010,
(Gram-positive Moderate (LE., (Inhibition zone) 2012a)
bacteria) LH.) Microwell
dilution assay (MIC)
Enterobacter Moderate (L.) Disc diffusion method Lazreg-Aref et al. (2012a)
cloacae (Inhibition zone)
(Gram-negative
bacteria)
Escherichia coli Moderate (LC., Paper diffusion assay Lazreg Aref et al. (2010,
(Gram-negative LE., LH.) (Inhibition zone) 2012a) and Rashid et al.
bacteria) Strong (L.) Well diffusion method (2014)
(Inhibition zone)
Microwell dilution
assay (MIC)
Klebsiella Strong (L.) Well diffusion method Rashid et al. (2014) and
pneumonae (Inhibition zone) Ahmed (2016)
(Gram-negative
bacteria)
Klebsiella spp. Strong (L.) Well diffusion method Ahmed (2016)
(Gram-negative (Inhibition zone)
bacteria)
Proteus mirabilis Strong (L., LC.,Paper diffusion assay Lazreg Aref et al. (2010,
(Gram-negative LE., LH.) (Inhibition zone) 2011b)
bacteria) Disc diffusion method
(Inhibition zone)
Microwell dilution
assay (MIC)
Pseudomonas Weak (LC., LE.) Paper diffusion assay Lazreg Aref et al. (2010,
aeruginosa Moderate (LH.) (Inhibition zone) 2012a) and Rashid et al.
(Gram-negative Strong (L.) Disc diffusion method (2014)
bacteria) (Inhibition zone)
Well diffusion method
(Inhibition zone)
Microwell dilution
assay (MIC)
(continued)
628 M. M. Hegazy et al.

Table 27.5 (continued)

Microorganism and Activities of
type different extracts Methodology Reference
Salmonella spp Strong (L.) Well diffusion method Ahmed (2016)
(Gram-negative (Inhibition zone)
bacteria)
Salmonella typhi Strong (L.) Well diffusion method Rashid et al. (2014)
(Gram-negative (Inhibition zone)
bacteria)
Seratia spp Moderate (L.) Well diffusion method Ahmed (2016)
(Gram-negative (Inhibition zone)
bacteria)
Staphylococcus Strong (L., LH.) Paper diffusion assay Lazreg Aref et al. (2010,
aureus Moderate (LC.) (Inhibition zone) 2012a) and Rashid et al.
(Gram-positive Disc diffusion assay (2014)
bacteria) (Inhibition zone)
Well diffusion method
(Inhibition zone)
Microwell dilution
assay (MIC)
Staphylococcus Moderate (L.) Paper diffusion method Lazreg-Aref et al. (2012a)
epidermidis Strong (LH.) (Inhibition zone) and Al-Sokari and El Sheikha
(Gram-positive Disc diffusion method (2015)
bacteria) (Inhibition zone)
Staphylococcus Strong (LH.) Disc diffusion method Lazreg-Aref et al. (2012a)
saprophyticus (Inhibition zone)
(Gram-positive
bacteria)
Streptococcus Strong (L.) Well diffusion method Rashid et al. (2014)
pyogenes (Inhibition zone)
(Gram-positive
bacteria)
L Latex, LH hexanoic fraction, LE Ethyl acetate fraction, LC chloroformic fraction

polar ones, and so both extracts showed the highest activities (Lazreg Aref et al.,
2010). These kinds of differences in susceptibility among microorganisms against
antimicrobial substances in F. carica latex different extracts may be explained by
the differences in cell well composition and or inheritance genes on plasmids that
can be easily transferred among bacterial strains or even due to the existence of dif-
ferent plant varieties (Lazreg Aref et al., 2010, 2011b). Finally, a great interest in
acquiring more knowledge regarding the antimicrobial effects ascribed to caprifig
latex exists among the scientific community since there is increasing evidence from
prospective observational studies that caprifig latex lowers the risk of infectious
diseases along with being effective for the treatment of superficial fungal and bacte-
rial infections, particularly in cutaneous mycosis. Moreover, another interesting
potential application of F. carica against fungus-induced diseases of crops results in
substituting phytotoxic and environmentally harmful substances with non-toxic
caprifig latex. Latex could also be used as a natural food preservative in the food
27 Composition and Biological Activities of Ficus carica Latex 629

Table 27.6 Antifungal activities of F. carica latex different fractions

Activities of
Microorganism and different
type extracts Methodology Reference
Aspergellus Weak (L., LH.)
Agar incorporation Lazreg Aref et al. (2010)
fumigatus Moderate (LC.)
(Percentage of
(Hyphamycets yeast) Strong (LE.) inhibition)
Aspergillus nigar Strong (L.) Well diffusion method Rashid et al. (2014)
(Fungi) (Inhibition zone)
Candida albicans Weak (LH.) Agar incorporation Lazreg Aref et al. (2010)
(Opportunists Strong (L., LC., (Percentage of and Rashid et al. (2014)
pathogenic yeast) LE.) inhibition)
Well diffusion method
(Inhibition zone)
Candida cruzi Moderate (L.) Well diffusion method Ahmed (2016)
(Yeast) (Inhibition zone)
Candida glabrata Weak (LE.) Agar incorporation Lazreg Aref et al. (2011b)
(Yeast) Moderate (L.) (Percentage of
Strong (LC.) inhibition)
Wells punched method
(Inhibition zone)
Candida kefyr Moderate (L.) Well diffusion method Ahmed (2016)
(Yeast) (Inhibition zone)
Candida sojae Strong (L.) Well diffusion method Ahmed (2016)
(Yeast) (Inhibition zone)
Candida tropicalis Strong (L.) Well diffusion method Ahmed (2016)
(Yeast) (Inhibition zone)
Cryptococcus Weak (LH.) Well diffusion method Lazreg Aref et al. (2010)
neoformans Moderate (L.) (Inhibition zone) and Ahmed (2016)
(Opportunists Strong (LC., Agar incorporation
pathogenic yeast) LE.) (Percentage of
inhibition)
Fusarium oxysporum Strong (L.) Well diffusion method Rashid et al. (2014)
(Fungi) (Inhibition zone)
Microsporum canis Moderate (LC.) Agar incorporation Lazreg Aref et al. (2010)
(Dermatophytes Strong (L., LE., (Percentage of and Al-Sokari and El
yeast) LH.) inhibition) Sheikha (2015)
Paper diffusion method
(Inhibition zone)
Trichophyton rubrum Weak (L., LC., Agar incorporation Lazreg Aref et al. (2010)
(Dermatophytes LH.) (Percentage of
yeast) Moderate (LE.) inhibition)
Trichophyton Weak (L., LC., Agar incorporation Lazreg Aref et al. (2010)
soudanense LE., LH.) (Percentage of
(Dermatophytes inhibition)
yeast)
L Latex, LH hexanoic fraction, LE Ethyl acetate fraction, LC chloroformic fraction
630 M. M. Hegazy et al.

processing industry, especially when we consider that fig latex has a high content of
minerals, vitamins, antioxidants, fiber, and proteins, which makes it valuable func-
tional food (Raskovic et al., 2016).

3.2 Antiviral Activity

The fig tree latex could be helpful in the treatment of bacterial and viral infections
caused by strains resistant to conventional drugs (Lazreg Aref et al., 2011a). Extracts
of F. carica latex have been found to inhibit the replication of Herpes simplex type
1 (HSV-1), Echovirus type 11 (ECV-11), and adenovirus (ADV). Moreover, they
have been found to act against several drug-resistant pathogens; among compounds
isolated from F. carica, it is found that lupeol (46), α-amyrin (43), and luteolin (12)
could be used as promising inhibitors against SARS-CoV-2 main protease which
plays a dynamic role in mediating viral replication and transcription. The molecular
dynamics simulation study revealed that among all the phytochemicals, α-amyrin
(43) is the most stable phytochemical, which could inhibit SARS-CoV-2 main pro-
tease strongly. The binding free energy analysis shows that α-amyrin (43) and lupeol
(46) have higher binding free energy in contrast to the known SARS-CoV-2 Mpro
inhibitor α-ketoamide (Ali et al., 2020). The antiviral activity of F. carica latex is
statistically significantly against Herpes simplex type 2 (HSV-2) lower than that of
acyclovir. In particular, the synergy produced by the antiviral activity of F. carica
and acyclovir combined had a more promising effect against HSV-2 than acyclovir
alone. It has been determined that the antiviral activity of F. carica is increased due
to the polysaccharides. The activity of F. carica latex against HSV-2 was confirmed
by a significant decrease in the number of viral copies (Ay & Duran, 2018). The fig
tree latex application on pox lesions showed continuous regression and shrinking of
the nodules from the fourth day of treatment (Abid & Ali, 2014). The hexanic and
hexane-ethyl acetate extracts inhibited the multiplication of HSV-1, ECV-11, and
ADV at 78 mg/ml concentrations. These two extracts are the most potent candidates
for herbal medicines to treat viral infectious diseases such as herpes virus, echovi-
rus, and adenovirus (Badgujar et al., 2014). Fig tree latex had a short duration ther-
apy to treat warts (Verruca vulgaris), flanking, and shrinking of responsive warts
after initiation of fig tree latex therapy (Bohlooli et al., 2007).

3.3 Anticancer Activity

Earlier studies have reported the antiproliferative effects of F. carica latex against
cells derived from esophageal and stomach cancers separately. The overall knowl-
edge suggests that fig latex could decrease tumor growth without adversely affect-
ing hematological and histological factors (Ghandehari & Fatemi, 2018; AlGhalban
et al., 2021). Latex of F. carica uses the different molecular mechanisms of action,
27 Composition and Biological Activities of Ficus carica Latex 631

including antiproliferative and anti-metastatic effects, along with significant effects

on cell shape and fighting against side effects of oxidative stress of free radicals
generated by the immune system cells (Ghandehari & Fatemi, 2018). Activities
were expected due to phytochemicals like 6-O-acyl-β-D-glucosyl-β-sitosterol and
its palmitoyl derivatives and antioxidant compounds, including flavonoids and poly-
phenolic compounds. Proteolytic enzymes abundantly accumulated in fig’s latex are
the major antiproliferative agents explaining the anticancer potential (Mavlonov
et al., 2008; Hashemi et al., 2011; Raskovic et al., 2016). The phenolic compounds
in fig latex are the main inhibitors of tumor growth, like protocatechuic acid derived
from latex which is partially responsible for the induction of cell death and inhibi-
tion of invasion in these cell lines. The fig latex possesses a group of proteases such
as ficin, caseinolytic, and gelatinolytic enzymes that induce apoptosis in cancer
cells without causing any side effects on normal cells (Ghandehari & Fatemi, 2018;
AlGhalban et al., 2021). Interestingly, the fig latex could inhibit the proliferation of
cancer cell lines without any cytotoxic effect on normal human cells with several
mechanisms (Hashemi et al., 2011). The antiproliferative actions were described
regarding fig latex; mainly its potential for apoptosis induction via interaction with
cell cycle causing rises of G0/G1phase cell population and decreases in S and G2/M
phase cell population (Ghandehari & Fatemi, 2018). Another supposed mechanism
for such effects might be associated with the inhibition of DNA synthesis, apoptosis
induction, cell cycle arrest of cancer cells, and extrinsic death receptor and intrinsic
mitochondrial-dependent apoptotic signaling pathway (Wang et al., 2008; Shin
et al., 2017). Given the fact that angiogenesis and inflammations are playmakers in
cancer progression, therefore the inhibitory effects of both factors by fig latex are a
major cytotoxicity mechanism. As known, platelets protected cancer cells from the
immune system, and they exacerbated metastasis by producing fibrin that caused
connection of cells to the endothelium of vessels. Therefore, fig latex preventable
effect of platelets increases in the number and/or activity further confirm its anti-
angiogenesis and anti-inflammatory potentials. Although mean platelet volume
(MPV) and platelet distribution width (PDW) are early indices of platelet activation,
while no significant difference was observed in the MPV, PDW and/or platelet
(PLT) upon administration of fig latex (Ghandehari & Fatemi, 2018). The anti-
angiogenesis and anti-inflammatory effects also were confirmed through a signifi-
cant decrease in the production of tumor necrosis factor-alpha, prostaglandin E2 and
vascular endothelial growth factor (Ghandehari & Fatemi, 2018). Fig latex also pre-
vents invasion through induction of lethal-7d (let-7d) expression (Tezcan et al.,
2015). Moreover, fig latex inhibited the proliferation of endothelial cells of central
vein cultured in a three-dimensional medium with collagen matrix and suppressed
the production of capillary tubules from these cells without damaging them
(Ghandehari & Fatemi, 2018). Interestingly, fig latex showed preventative potential
against cancer induced anemia which is the most common side effects caused by
neoplastic diseases via inhibition of erythropoietin secretion. The anti-inflammatory
effect of the fig latex prevented the destruction of erythropoietin-secreting tissues
and, accordingly, prevented a decrease in RBC and blood cell indices (Ghandehari
& Fatemi, 2018). Furthermore, consistent changes in cell shape after treatment with
632 M. M. Hegazy et al.

the latex were observed by microscopical analysis. The cells showed morphological
changes such as round shape and shrinkage, which indicated a stoppage of prolif-
eration and turning on apoptosis. Although a round shape was the dominant mor-
phology, other cell shapes like cell blebbing, cytoplasmic vacuolation, and uneven
cell structure and distribution confirmed apoptosis (AlGhalban et al., 2021).
Reduced expression of phosphorylated map kinase (ERK2), cAMP-response
element-binding protein (CREB), and glycogen synthase kinase-3a/b (GSK-3a/b)
with fig latex treatment which is actively involved in cellular proliferation processes
(AlGhalban et al., 2021). To evaluate the cytotoxic activities of fig latex, different
methodologies were applied. Examples include DNA damage assay, MTT assay,
wound healing assay, tumors mean volumes measurement, WST-1 assay, chick cho-
rioallantoic membrane assay, and morphological studies (Ghandehari & Fatemi,
2018; AlGhalban et al., 2021). Antiproliferative activities were reported in human
colon cancer cell line (HT-29) (Soltana et al., 2019), human glioma (HCC) (Wang
et al., 2008), human oral cancer cells (Shin et al., 2017), glioblastoma multiforme
(GBM) (Tezcan et al., 2015), stomach cancer (Hashemi et al., 2011), Melanoma
cells (A375) (Menichini et al., 2012), human hepatocellular carcinoma (HepG˗2),
human breast cancer cells (MCF˗7), and human colon cancer cell line (HCT˗116)
(Yahiaoui et al., 2022). In conclusion, the administration of fig latex prevented the
oversized growth of cancerous tumors without causing side effects on hematologic
parameters and induction of the inflammation in tissues that can be attributed to
antioxidant, anti-inflammatory, and pro-apoptotic effects this natural product.

3.4 Anti-Angiogenic Activity

Angiogenesis, a complex multi-step process resulting in new blood vessels from

preexisting ones, represents a critical process for oxygen and nutrient supply to
proliferating cells. Therefore, promoting tumor growth and metastasis. Inhibition of
tumor angiogenesis is therefore considered a promising anticancer target. It was
reported that latex extracts of F. carica possess strong antiangiogenic and anti-
proliferative activities (Mostafaei et al., 2011). Latex from immature fruits of sev-
eral different F. carica varieties were tested for anti-angiogenic activity in a unique
system involving the use of segments of placental veins and human tumor biopsies.
Three out of six of these varieties demonstrated significant anti-angiogenic activity
yet were non-toxic. This non-toxic, anti-angiogenic activity was proved using a
human umbilical vein endothelial cell (HUVEC) tubule formation assay and MTT
antiproliferation assay (Pawlus et al., 2008).
Bioactive compounds isolated from F. carica latex-like 6-O-acyl-β-D-glucosyl-
β-sitosterol derivatives, i.e., AGS (acyl moiety: palmitoyl, linoleoyl, stearyl, and
oleyl). Palmitoyl derivative of AGS acts as the most active inhibitor for various
cancer cell lines compared with other derivatives. For example, AGS is reported for
in vitro inhibition of Burkitt lymphoma cell line (DG-75) and prostatic cancer cell
lines (DU-145). Thus, AGS is the most influential anticancer agent. Therefore, latex
27 Composition and Biological Activities of Ficus carica Latex 633

extract could be an ideal candidate as a prospective agent for preventing angiogen-

esis in cancer and other chronic disorders (Badgujar et al., 2014).
Furthermore, fig latex can prevent the diffusion of cancer cells into new areas and
the formation of metastatic tumors. Treatment with fig latex has allowed the cell
cycle exit by inducing the cell cycle arrest in the Sub G1 phase in cervical cancer
cell lines. This finding demonstrates the ability of fig latex to prevent cell progres-
sion and proliferation. Cell cycle deregulation is known to be a significant element
in cervical cancer carcinogenesis hence regulation of cancer cells division through
induction of apoptosis is an efficient anti-cancer strategy (Ghanbari et al., 2019).
Latex of F. carica inhibited the proliferation of cancer cell lines but did not indicate
any cytotoxic activity against normal cells in vitro. Latex of F. carica had a promis-
ing effect in inhibiting stomach cancer cell line growth but without any clear effect
on peripheral blood mononuclear cells. Ficin, a cysteine proteinase isolated from
the latex of F. carica tree. Cysteine proteinases are a group of enzymes leading to
the apoptosis of cancer cells. Furthermore, the anticancer effects may be associated
with antioxidant properties due to its phenolic components. Additionally, We con-
cluded that F. carica has an anticancer cell activity while cancer cells were more
sensitive to this latex than normal cells (Hashemi et al., 2011).

3.5 Antioxidant Activity

Antioxidant compounds, such as phenolics, and organic acids, protect against oxi-
dation or cellular damage caused by reactive species, preventing the initiation of
several diseases, like many types of cancer, heart disease, diabetes, and neurodegen-
erative illnesses (Garcı́a-Alonso et al., 2004). The phenolic content of latex extracted
from the fruits of F. carica, collected from Ministry of Agriculture, Sharkia govern-
ment, Egypt, was effective in scavenging free radicals of DPPH (2,2-diphenyl-1-
picrylhydrazyl), and ABTS (2,2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic
acid)), with IC50 values of 13.6, and 4.5, μg GAE/ml, respectively (Abdel-Aty et al.,
2019). Its total antioxidant activity, using phosphomolybdenum assay, had been
reported with EC50 values of 39 μg GAE/mL (Abdel-Aty et al., 2019). The antioxi-
dant ability of latex extracted from immature green fruits of F. carica, cultivar pingo
de Mel trees growing in the Mirandela region (Northeast Portugal), was screened by
DPPḤ assay, which allowed observation of a concentration-dependent potential
(IC25 = 1049 μg/mL) (Oliveira et al., 2010b). The analyzed sample presented a pro-
tective effect against superoxide radical, in a concentration-dependent way, with an
IC25 at 291 μg/mL (Oliveira et al., 2010b). It displayed nitric oxide scavenging
capacity, which was concentration-dependent (IC25 = 1768 μg/mL) (Oliveira et al.,
2010b). Polyphenols and flavonoids extracted from the latex collected from the
neck of F. carica L. fruit before ripening from Erbil city, north of Iraq, showed a
significant reduction in the rate of superoxide dismutase and glutathione reductase
enzymes in liver cells of mature male Wistar albino rats (Rattus norvegicus) (Aziz,
2012). Ficin had a DPPḤ radical scavenging ability of about 78.7% compared to
634 M. M. Hegazy et al.

L-ascorbic acid at the highest concentration of 1000 μg/mL. In addition, ficin had
an H2O2 scavenging ability of about 50.5% compared to L-ascorbic acid at the high-
est concentration of 1000 μg/mL. These results suggest that Polyphenols, flavo-
noids, and ficin in the latex of F. carica are responsible for its anti-oxidant activity
(Aziz, 2012; Cho et al., 2019).

3.6 Anti-Inflammatory Activity

Ficus carica has been exhibited as an anti-inflammatory remedy in traditional medi-

cine (Werbach, 1993; Duke, 2002; Mawa et al., 2013). Experimentally, it was found
that NO (nitric oxide) production was significantly decreased by ficin in a
concentration-dependent manner: at a concentration of 10, 25, 50, and 100 μg/mL,
NO production was significantly reduced to 90.5%, 77.3, 67.9, and 59.6%, respec-
tively, compared to that in the lipopolysaccharide (LPS) treated group without ficin
treatment (Cho et al., 2019). As well as ficin could decrease the amount of phos-
phorylated IκB and Nuclear factor κB (NF-κB) protein induced by LPS in a
concentration-dependent manner (Cho et al., 2019).

3.7 Anticoagulant Activity

Latices of several Ficus species contain ficin. Ficin (a single polypeptide chain
active at neutral pH while inactive at pH below 3.0) is a heterogeneous group of
proteases. The procoagulant activity of F. carica crude extract could be explained
by cysteine proteases FPI, FPII, and serine protease FPIII, which are types of ficin.
This procoagulants characteristics of cysteine proteases might be due to their ability
to cleave the Aα and Bβ chains, but not the γ chain of fibrinogen, releasing fibrino-
peptide A and B, respectively, forming fibrin clot that is soft and friable or their
ability to activate human factor X by cleave it to create FXa. However, the antico-
agulant activity of F. carica crude extract might be due to the complete digestion of
fibrinogen or the presence of serine protease inhibitor that inhibited coagulation
factors or antiplatelet effect so that F. carica latex has procoagulant and anticoagu-
lant properties at the same time. Latex of F. carica can control its prominent coagu-
lation stimulant nature caused by ficin (more than 5 isoenzymes) by fibrinolytic
serine proteases. The anticoagulant property of the latex at a higher protein concen-
tration suggested the presence of other anti-proteases and small peptides that may
inhibit ficin (Hamed et al., 2020). Ficin shortened the activated partial thromboplas-
tin time and the prothrombin time of normal plasmas and plasmas deficient in coag-
ulation factors; this showed that the hemostatic potency of Ficus proteases was
based on the activation of human coagulation factor X (FX), which could explain
the use of these latices as a local hemostatic agent in natural medicine (Richter
et al., 2002).
27 Composition and Biological Activities of Ficus carica Latex 635

3.8 Antihypertensive Activity (Angiotensin I-Converting

Enzyme Inhibitors Activity)

Angiotensin I-converting enzyme catalyzes the production of the vasoconstrictor

angiotensin II and the inactivation of the vasodilator bradykinin. Three peptides
were isolated from the latex of Ficus carica; sequences of these peptides were: Ala-
Val-Asn-Pro-Ile-Arg (FLP-1), Leu-Tyr-Pro-Val-Lys (FLP-2), and Leu-Val-Arg
(FLP-3) are compared with ACE isolated from rabbit lung Hippuryl-L-histidyl-L-
leucine (Hip-His-Leu) with IC50 13, 4.5 and 14 μM, respectively. They revealed the
Angiotensin I-converting enzyme inhibitor effect by catalyzing His-Leu release
from the carboxyl-terminal angiotensin. Therefore, ACE inhibitors derived from
F. carica latex may be categorized along with natural peptide inhibitors (Maruyama
et al., 1989).

3.9 Hepatoprotective Activity

The extract of latex, collected from the neck of F. carica L. fruit before ripening,
from Erbil city, north of Iraq, showed attenuated lead hepatotoxicity in rats; through
one or more of these suggested mechanisms of action; even by lowering the oxida-
tive stress, second: increasing the oxidant enzymes level, and third: acting as a che-
lating agent for lead ions. In addition, it was evaluated by induction of oxidative
stress using lead acetate in rats. The F. carica latex was highly successful in attenu-
ating lead hepatotoxicity due to its high total phenol and flavonoid contents
(Aziz, 2012).

3.10 Antiparasitic Activity

Latex of F. carica collected from the garden of Kufa University, Iraq, in a compara-
tive study with sodium stibogluconate, showed promising antileishmanial activity
in a concentration of 500 μg/mL. Complete healing in the ninth week was (85.9%)
and without side effects on the Leishmania tropica parasite (Shlash et al., 2021). On
the other hand, the fig latex had anthelmintic activity (de Amorin et al., 1999), espe-
cially against Ascaris and Tricharus. This effect has been attributed to the presence
of ficin in fig latex (an enzyme that has been recognized for its anthelmintic activity)
(Joseph & Raj, 2011).
636 M. M. Hegazy et al.

3.11 Adverse Reactions

Although the fruit of F. carica is edible, it is rarely described as a cause of food

allergy. The latex contains ficin, a proteolytic enzyme that causes pruritus (Arthur &
Shelley, 1955) and acts as an irritant on inflamed skin (Budavari et al., 1996). Latex
found in the leaves and stems seems to contain irritant and phototoxic chemicals,
although one case report suggests that the fruit also contains photosensitizers (Ippen,
1982). Skin contact with latex may produce allergic reactions like dermatitis,
asthma, and anaphylaxis (Bohlooli et al., 2007).

4 Conclusion

Several studies have investigated fig latex bioactive components and pharmacologi-
cal and industrial activities. In conclusion, the administration of fig latex provides
multitask curative features without causing side effects attributed to its phytochemi-
cal constituents. On the other hand, oxidative stress, inflammation, viral infection,
microbial infection, hepatotoxicity, hypertension, parasitic invasion, blood coagula-
tion, angiogenesis, and oversized growth of cancerous tumors inhibition were
reported for fig latex. Furthermore, the phenolic content is responsible for the most
pharmacological activities, specially mangiferin potentials as antidiabetic, antimi-
crobial, anticancer antioxidant phytoconstituent. Moreover, fig latex is considered a
rich source of proteases such as ficin, caseinolytic, and gelatinolytic enzymes
involved in several activities, including anticancer without side effects on normal
cells and as a promising alternative to animal enzymes in milk clotting. Finally, fig
latex needs further standardization procedures as a pharmaceutical product.

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Chapter 28
Extraction and Analysis of Polyphenolic
Compounds in Ficus carica L.

Babra Moyo and Nikita T. Tavengwa

1 Introduction

Ficus carica L, a plant that belongs to the Moraceae family, is one of the most
widely farmed plants in the world because of its richness in phytochemicals that can
have potential applications in pharmaceutical, cosmetic, and food industries
(Amessis-Ouchemoukh et al., 2017; Khadhraoui et al., 2019; Maghsoudlou et al.,
2017). This species is distinguished by a variety of chemical components which
have been beneficial to human health for millennia (Ammar et al., 2015; Bachir Bey
et al., 2017; Mopuri et al., 2018). Figs are mainly recognized for their medicinal
characteristics, such as their usage in traditional medicine to treat cardiovascular
(Alamgeer et al., 2017) and anti-inflammatory problems (Allahyari et al., 2014;
Amessis-Ouchemoukh et al., 2017).
Phenolics, including phenolic acids and flavonoids, are responsible for the health
benefits of these compounds found in various fruits and vegetables, including figs
(Arvaniti et al., 2019; Nakilcioğlu-Taş & Ötleş, 2021). Interest in extracting pheno-
lic compounds from plant materials such as fruits and vegetables has grown in the
past years as demand for natural antioxidants increases. Therefore, there is a need to
develop efficient extraction methods that can be used to isolate these compounds
from plant materials.

B. Moyo
Department of Biochemistry and Microbiology, Faculty of Science,
Engineering and Agriculture, University of Venda, Thohoyandou, South Africa
Department of Food Science and Technology, Faculty of Science,
Engineering and Agriculture, University of Venda, Thohoyandou, South Africa
e-mail: [email protected]
N. T. Tavengwa (*)
Department of Chemistry, Faculty of Science, Engineering and Agriculture,
University of Venda, Thohoyandou, South Africa
e-mail: [email protected]
© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 643
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_28
644 B. Moyo and N. T. Tavengwa

The structural diversity of secondary metabolites coupled with the high com-
plexity of the plant metabolome poses a challenge in selecting a suitable extraction
technique for the pre-concentration of these compounds. This implies that an inef-
ficient extraction or a single-step method might influence the isolation of these com-
pounds from the plant samples. As a result, it is critical to select a suitable extraction
technique to isolate structurally diverse phytocompounds (Alara et al., 2021).
Solvent extraction is the commonly used technique for extracting fig crude extracts
(Ersoy et al., 2017; Konak et al., 2017; Meziant et al., 2018; Pereira et al., 2017).
Selecting a suitable solvent for this extraction technique is of uttermost importance
since the amount and the nature of phytochemicals extracted will depend on the
extraction solvent. Therefore, the physicochemical properties of the analytes of
interest and the analysis instrument should be considered (Choi et al., 2021; Kang
et al., 2019). Other methods that have been applied in the past years for the extrac-
tion of various polyphenols from various plant samples include pressurized liquid
extraction (Barrales et al., 2018; Benito-Román et al., 2015), ultrasound-assisted
extraction (Bakirtzi et al., 2016; Bosiljkov et al., 2017; Um et al., 2018) and
microwave-assisted extraction (Alara et al., 2018; Sarfarazi et al., 2020). However,
the application of these extraction techniques in figs is still limited.
Although few reviews have been published on the extraction of polyphenols from
figs, to the best of our knowledge, none have reviewed extraction methods in detail.
Arvaniti et al. (2019) reviewed the extraction of phytocompounds from figs using dif-
ferent techniques such as solid-liquid extraction (SLE), solid-phase extraction, and
subcritical water extraction, to mention a few. However, the principles of these meth-
ods were not discussed in depth. In a review authored by Solana and Romano (2019),
the analysis of polyphenols in beverages produced from figs using colorimetric
assays. However, in this review, extraction methods were not discussed in detail,
given that sample preparation is a paramount step in analyzing phytochemicals. In
this context, the application of conventional SLE in extracting polyphenols from fig
fruits was discussed in this chapter. In addition, non-conventional extraction based on
SLE, such as subcritical water extraction and microwave-assisted extraction, were
discussed briefly, given that their application in this particular matrix is still limited.

1.1 Nutritional and Health Benefits of Ficus carica L.

The fig (Ficus carica L.) is one of the earliest fruit trees cultivated globally (Ersoy
et al., 2017). It is a popular crop worldwide and is consumed widely in Mediterranean
diets (Alexandre et al., 2017; Barolo et al., 2014; Harzallah et al., 2016). The differ-
ent parts of the fig plant are used for food and therapeutic purposes (Ammar et al.,
2015; Debib et al., 2016). Fig fruits have long been used in traditional medicine for
their laxative, pulmonary, antispasmodic, and anti-inflammatory properties (Barolo
et al., 2014; Nadeem & Zeb, 2018; Rodríguez-Solana et al., 2018; Viuda-Martos
et al., 2015). These health-promoting benefits are primarily because figs contain
significant levels of polyphenols, which have antioxidant properties and can help
prevent diseases caused by oxidative stress (Amessis-Ouchemoukh et al., 2017;
28 Extraction and Analysis of Polyphenolic Compounds in Ficus carica L. 645

Bahrin et al., 2018; Harzallah et al., 2016; Nakilcioğlu-Taş & Ötleş, 2021). These
compounds help reduce cellular damage caused by reactive species such as hydro-
gen peroxide while acting as scavengers for free radicals produced by oxygen or
nitrogen (Amessis-Ouchemoukh et al., 2017; Solana & Romano, 2019). Moreover,
figs are high in crude fiber, minerals, and vitamins, and they are lipid, sodium, and
cholesterol free (Buenrostro-Figueroa et al., 2017; Nakilcioğlu-Taş & Ötleş, 2021).
The pharmaceutical industry is increasingly becoming interested in natural products
to produce drugs as alternatives to synthetic drugs (Barolo et al., 2014). For this reason,
the therapeutic qualities of the genus Ficus have been widely explored during the last
few decades through biological activity tests. Antibacterial (Benmaghnia et al., 2021;
Palmeira et al., 2019), antidiabetic (Mopuri et al., 2018; Wojdyło et al., 2016), anticon-
vulsant (Raafat & Wurglics, 2019), and hepatoprotection (Debib et al., 2016) properties
are some of the biological activities that have been reported for Ficus carica L. The
by-products resulting from fig processing, such as the fig skin, have been reported to
possess high quantities of polyphenol compounds (Cheynier et al., 2013; Harzallah
et al., 2016; Kamiloglu & Capanoglu, 2015). Dark-colored figs have been reported to
have high polyphenols due to their higher anthocyanin content (Kamiloglu & Capanoglu,
2015; Maghsoudlou et al., 2017; Meziant et al., 2018). Therefore, it is advisable to con-
sume the whole fruit as the skin has more phenolic compounds. Fig seeds are also
thought to play a significant role in the nutrient content and health benefits of the fruit
(Nakilcioğlu-Taş & Ötleş, 2021).

1.2 Plant Secondary Metabolites

Secondary metabolites are a diverse group of naturally occurring compounds syn-

thesized by plants for their reproduction and defense against biotic (viruses, bacte-
ria, fungi, and vertebrates) and abiotic stress (UV light, drought, contamination with
heavy metals, and temperature changes) (Khare et al., 2020; Tungmunnithum et al.,
2018). These compounds have a high potential for use as drugs by public healthcare
systems (Boy et al., 2018; Choi et al., 2021). Secondary metabolites are classified
according to their biosynthetic pathway, chemical structure, composition, or solu-
bility (Soukup & Soukup, 2015; Thirumurugan et al., 2018). Alkaloids, terpenes,
and phenolics are the major compounds produced by plants (Kabera et al., 2014;
Katz & Baltz, 2016; Alamgeer et al., 2017). However, only polyphenols will be
discussed in this paper as they are widely studied in fig fruits.

1.2.1 Phenolics

Secondary metabolites known as phenolics are structurally diverse compounds that

are found in most plant species, including Ficus carica L. (Ammar et al., 2015;
Nakilcioğlu-Taş & Ötleş, 2021). They can exist as aglycones, glycosides, polymer-
ized or monomer structures, and matrix or free-bound molecules (Alara et al.,
2021). Phenolics are the most abundant group of secondary metabolites found in
646 B. Moyo and N. T. Tavengwa

figs, and they are responsible for the color, taste, and flavor of these fruits (Ammar
et al., 2015; Garagounis et al., 2021; Vaishnav & Demain, 2011). The presence of
one or more phenol groups distinguishes them, and they range in complexity from
basic molecules with one aromatic ring to highly complex polymeric compounds
(Pandey & Rizvi, 2009). Some phenolics found in figs, like quercetin and narin-
genin, are pharmacologically essential compounds because of their anti-
inflammatory properties (Cheng et al., 2019; Karuppagounder et al., 2016).
Genistein, also found in figs, possesses phytoestrogenic activity (Farmer et al.,
2016; Morales-Otal et al., 2016). In addition, various phenolic compounds, like
flavonoids, are potent antioxidants because of their free radical scavenging effect
(Panche et al., 2016; Rodríguez-García et al., 2019). Phenolics are classified accord-
ing to their biosynthetic pathway or structure into simple phenolic acids, flavonoids,
tannins, coumarins, lignans, and stilbenes, depending on the number of phenol rings
that they contain and the moieties that bind these rings (Abegaz & Kinfe, 2019;
Hussein & El-Anssary, 2019). The two types of phenolic acids are hydroxycinnamic
and hydroxybenzoic acids, and these have been reported in figs (Ammar et al.,
2015; Nakilcioğlu-Taş & Ötleş, 2021). An example of simple benzoyl phenolic acid
is gallic acid, a parent compound of gallotannins, whereas p-coumaric, caffeic, and
ferulic acids are examples of hydroxycinnamic acids (Saibabu et al., 2015). Ammar
et al. (2015) reported the presence of simple phenolic acids in figs, including gallic,
vanillic, caffeic, and ferulic acids.
Chalcones, flavans, flavanols, flavanone, flavone, isoflavone, anthocyanidins, and
xanthones are the different types of flavonoids. Hundreds of compounds in each group
vary in the number and positioning of hydroxyl groups on the flavan scaffold (Abegaz
& Kinfe, 2019). Anthocyanins are responsible for the coloration of figs, and these
compounds can be used as food colorants (Ammar et al., 2015; Backes et al., 2018).
Quercetin is a flavonol that is one of nature’s most effective antioxidants and a key
component in figs (Maghsoudlou et al., 2017; Wojdyło et al., 2016). Tannins exist as
hydrolyzable tannins and condensed tannins. Hydrolyzable tannins consist of several
molecules of phenolic acids, such as gallic and hexahydroxydiphenic acids, attached
to a central glucose molecule through an ester bond (Durazzo et al., 2019).
On the other hand, condensed tannins, or proanthocyanidins, are compounds
with structures based on oligomeric flavonoid precursors that differ in the type of
linkage between flavonoid units (Fraga-Corral et al., 2020). However, few studies
have investigated the presence of tannins in figs (Alexandre et al., 2017). Coumarins
are derivatives of benzo-α-pyrone, which is an o-hydroxycinnamic acid lactone
(Camara et al., 2020). Examples of coumarins detected in fig leaves and fruits
include umbellifone, psoralen, and methoxypsolaren (Ammar et al., 2015; Qin
et al., 2015). Figure 28.1 shows the general chemical structures of the common
polyphenols found in figs.
28 Extraction and Analysis of Polyphenolic Compounds in Ficus carica L. 647

Fig. 28.1 Chemical structures of the different classes of polyphenols

1.3 Extraction Methods of Polyphenols from Figs

Extraction of polyphenols from figs is the first step in removing interferences and
isolating these compounds from this matrix (Hikmawanti et al., 2021). The down-
stream analysis only identifies metabolites previously extracted; therefore, develop-
ing a proper sample preparation technique is critical for obtaining accurate results
(Ibañez et al., 2012). Developing efficient sample preparation techniques in plant
metabolomics is particularly challenging due to the complexity of plant biological
systems, making it challenging to determine target analytes reliably (Mastellone
et al., 2021). Moreover, given their occurrence in low concentrations in plant matri-
ces, it is critical to adopt extraction methods that enable the efficient recovery of
these compounds from bulk matrices while also removing matrix interferences
(Câmara et al., 2020). In the last few decades, detection technologies have grown at
an unparalleled rate (Aron et al., 2020; Nothias et al., 2020; Stefanucci et al., 2020).
Phenomenal developments have been made in high-resolution analytical techniques,
such as chromatography and spectroscopy, making phytochemical analysis much
easier than before. Nevertheless, sample preparation is still a bottleneck in analyz-
ing these compounds (Moyo & Tavengwa, 2021a). Moreover, there is no widely
established method for recovering all phenolics or those belonging to a specific
class from plant materials, including figs. As a result, an optimized procedure for
recovering phenolic compounds from plant sources is required (Alara et al., 2021).
The most widely used traditional extraction methods, including liquid-liquid extrac-
tion (LLE), solid-liquid extraction (SLE), Soxhlet extraction, and hydro-distillation,
are time-consuming, labor-intensive, consume large amounts of organic solvents
648 B. Moyo and N. T. Tavengwa

and have low extraction efficiencies (Mastellone et al., 2021). Accordingly,

“greener” options for pre-treatment plant samples have emerged over the past years
to overcome the disadvantages of classical extraction methods (Sarfarazi et al.,
2020). These methods include pressurized liquid extraction (PLE) (Barrales et al.,
2018; Benito-Román et al., 2015), pressurized hot water extraction (PHWE)
(Cvetanović et al., 2018; Lachos-Perez et al., 2020), and microwave-assisted extrac-
tion (MAE) (Alara et al., 2018; Sarfarazi et al., 2020). However, applying these
methods to extract polyphenols from figs is still very limited (Maghsoudlou et al.,
2017; Masota et al., 2020).

1.3.1 Conventional Extraction Techniques

The Application of SLE in the Extraction of Phenolics from Figs

Solid-liquid extraction (SLE), a technique where analytes of interest are extracted

using a liquid from a solid (plant material), is based on the principles of osmosis and
diffusion (Naviglio et al., 2019). First, the solvent penetrates through the plant mate-
rial matrix, and analytes of interest (polyphenols) are released into the solvent
through osmosis (Zhang et al., 2018). Subsequently, polyphenols disperse into the
solvent via diffusion (Milescu et al., 2020). Figure 28.2 shows the general SLE pro-
cedure, including the most reported sample pre-treatment steps from studies reviewed
in this chapter. Although SLE has been known for a long time, it is still widely
employed to extract phytochemicals from various plant materials, including figs.
Notwithstanding the simplicity of the SLE technique, it has drawbacks such as the
consumption of large volumes of organic solvents, inadequate extraction efficiency,
chemical transformation of extracts, and lengthy extraction times (Alexandre et al.,
2017; Amessis-Ouchemoukh et al., 2017; Mopuri et al., 2018; Vernès et al., 2020).
Furthermore, the high concentration of free sugars and other matrices can challenge
analyzing phenolic compounds from figs using this technique (Arvaniti et al., 2019).
Despite the mentioned challenges associated with SLE, it is notable that it is still
the most widely utilized technique for extracting polyphenols from the whole fruit
as well as other portions of figs, such as the peel and pulp, as given in Table 28.1
(Alamgeer et al., 2017; Amessis-Ouchemoukh et al., 2017; Ammar et al., 2015;
Bachir Bey et al., 2017; Bachir Bey & Louaileche, 2015; Benmaghnia et al., 2021;
Debib et al., 2016; Ersoy et al., 2017; Harzallah et al., 2016; Hoxha et al., 2015;
Hoxha & Kongoli, 2016; Kamiloglu & Capanoglu, 2015; Khadhraoui et al., 2019;
Konak et al., 2017; Mahmoudi et al., 2018; Meziant et al., 2018; Mopuri et al., 2018;
Nakilcioğlu-Taş & Ötleş, 2021; Pereira et al., 2017; Viuda-Martos et al., 2015;
Wojdyło et al., 2016). Prior to the SLE procedure, sample pre-treatment steps that
have been reported in the studies reviewed in this chapter involve drying the figs
using different methods of drying, such as shade drying (Maghsoudlou et al., 2017),
freeze-drying (Bachir Bey et al., 2017; Wojdyło et al., 2016) and sun drying
(Kamiloglu & Capanoglu, 2015; Khadhraoui et al., 2019; Konak et al., 2017). Dried
fig samples were then ground to increase the contact area between the sample and
the solvent (Njila et al., 2017), facilitating the analytes’ mass transfer. In some
28 Extraction and Analysis of Polyphenolic Compounds in Ficus carica L. 649

Fig. 28.2 General SLE procedure

studies, fresh fig samples were homogenized with the solvent (Ersoy et al., 2017).
After extraction, samples were centrifuged to remove suspended solids (Alexandre
et al., 2017; Amessis-Ouchemoukh et al., 2017; Nakilcioğlu-Taş & Ötleş, 2021).
Table 28.1 summarizes the application of different extraction methods to isolate
polyphenols from figs. Even though there is no universally accepted optimal solvent
for polyphenol extraction, it is often assumed that solvents with higher polarity are
suitable due to the increased solubility of polyphenols in such solvents (Alara et al.,
2021). The varying physicochemical properties of polyphenols significantly impact
the solubility of these compounds in solvents. Methanol (Bachir Bey et al., 2017;
Harzallah et al., 2016; Mahmoudi et al., 2018), ethanol (Amessis-Ouchemoukh
et al., 2017; Konak et al., 2017; Mopuri et al., 2018), and acetone (Pereira et al.,
2017) are the most commonly used solvents for extracting polyphenols from figs.
Different compositions of ethanol (Alexandre et al., 2017) and methanol, particu-
larly 80% methanol (Ammar et al., 2015; Benmaghnia et al., 2021; Hoxha et al.,
2015; Khadhraoui et al., 2019; Viuda-Martos et al., 2015), were also widely used as
extraction solvents. The selection of the solvent is crucial for solvent extraction
methods. Nonetheless, few studies were reviewed in this chapter that investigated
the effect of different solvents in extracting polyphenols (Backes et al., 2018;
Meziant et al., 2018; Mopuri et al., 2018). Extraction methods can be improved by
optimizing parameters that may affect the extraction efficiency of the analytes of
interest. Acidified organic solvents have also been recommended to prevent the deg-
radation of polyphenols (Meziant et al., 2018). Some of the acids that have been
reported include formic acid (Kamiloglu & Capanoglu, 2015; Pereira et al., 2017),
hydrochloric acid (Pereira et al., 2017), ascorbic acid (Wojdyło et al., 2016),
Table 28.1 Application of different extraction methods in the analysis of polyphenols in figs
Polyphenols Extraction Extraction procedure
Matrix analyzed method Extraction solvent after solvent addition Method of detection Concentration References
Dried figs TPC and TFC SLE Ethanol Maceration and UV/Vis 2190 mg GAE Amessis-Ouchemoukh
samples were stirred spectrophotometry 100 g−1 (TPC) et al. (2017)
for 24 h at room 858 mg QE 100 g−1
temperature (TFC)
Fermented fig TPC, TFC and High 9% ethanol (TPC), High pressure assisted
UV/Vis – Alexandre et al.
by-products anthocyanidins pressure 47% ethanol (TFC) extraction spectrophotometry (TPC (2017)
assisted and TFC) and UHPLC-
extraction PDA (phenolic acids)
Fig seeds TPC, TFC and 5 SLE 50% methanol (TPC, The sample mixture UV/Vis 714 mg GAE kg−1 Nakilcioğlu-Taş and
individual TFC and was shaken at 50 °C spectrophotometry (TPC (TPC) Ötleş (2021)
polyphenols polyphenols) for 90 min in a shaking and TFC) 312 mg CE kg−1
water bath HPLC-DAD (individual (TFC)
polyphenols)
Fresh figs TPC SLE 50% acetone and Homogenization of UV/Vis 25.4–37.2 mg GAE Ersoy et al. (2017)
80% methanol fresh samples with the spectrophotometry g−1 (TPC)
solvent in a mixer
Fig juice TPC and TFC SLE Methanol Homogenization of the UV/Vis 50.5–74.1 mg GAE Harzallah et al. (2016)
samples with the spectrophotometry g−1(TPC)
solvent in a 11.3–
homogenizer 12.7 mg CE kg−1
(TFC)
Dried figs 14 phenolic SLE Methanol Ultrasonication for 1 h HPLC-FLD 0.13–73 mg 100 g−1 Bachir Bey et al.
acids and 8 45 min (2017)
flavonoids
Dried figs TPC and TFC SLE 80% methanol Extraction on an orbital UV/Vis 73.7–201 mg GAE Khadhraoui et al.
shaker for 1 h spectrophotometry and g−1 (TPC) (2019)
LC-ESI-MS 57.9–
(polyphenols) 112 mg CE kg−1
(TFC)
Dried figs TPC SLE Ethanol The sample was UV/Vis 55.5–574 mg GAE Konak et al. (2017)
homogenized for 4 min spectrophotometry g−1
Fig pulp and skin TPC and TFC SWE Water Extraction at 160 °C UV/Vis – Maghsoudlou et al.
and at a pressure of spectrophotometry (2017)
6.89 MPa for 30 min
Fig pulp, skin and 91 phenolic SLE 80% methanol Ultrasonication UHPLC-QTOF-MS 0.26–0.32 mg GAE Ammar et al. (2015)
leaves compounds (profiling) and UV/Vis 100 g−1 (TPC)
(profiling) and spectrophotometry
TPC (TPC)
Dried figs TPC SLE 80% methanol Maceration in the UV/Vis 700 mg GAE Debib et al. (2015)
containing 1% solvent at 4 °C for 16 h spectrophotometry 100 kg−1
trifluoracetic acid
Dried figs TPC and TFC SLE 80% methanol Ultrasonication at low UV/Vis 167–930 mg GAE Kamiloglu and
containing 0.1% temperatures for spectrophotometry 100 g−1 (TPC) Capanoglu (2015)
formic acid 15 min 7–234 mg CE
100 g−1 (TFC)
Dried fig peels Monomeric SLE 80% methanol Stirring at room UV/Vis 346 mg 100 g−1 Meziant et al. (2018)
anthocyanins containing 0.1% temperature on a spectrophotometry
citric acid magnetic stirrer for
180 min
Dried figs TPC and TFC SLE Ethanol Extraction in the UV/Vis 14.6 mg GAE g−1 Mopuri et al. (2018)
solvent for 48–72 h spectrophotometry (TPC)
81.6 mg QE g−1
(TFC)
Fig pulp and peel TPC and TFC SLE 80% ethanol Stirring at room UV/Vis 26.7–169 mg GAE Pereira et al. (2017)
containing 1% temperature on a spectrophotometry 100 g−1 (TPC)
hydrochloric acid magnetic stirrer for 81.6 mg QE 100 g−1
20 min (TFC)
(continued)
Table 28.1 (continued)
Fig pulp and peal TPC and TFC SLE 80% methanol The sample was shaken UV/Vis 1.92–5.76 mg GAE Viuda-Martos et al.
for 2 min, followed by spectrophotometry g−1 (TPC) (2015)
ultrasonication for 2 h 9.24–19.1 mg RE
at room temperature g−1 (TFC)
Dried figs 10 polyphenols SLE 30% methanol Ultrasonication for UHPLC-QTOF-MS 0.5–328 mg 100 g−1 Wojdyło et al. (2016)
acidified with 1% 15 min
ascorbic acid
Dried figs TPC SLE 80% methanol Vortex mixing for UV/Vis 45.2–160 mg GAE Hoxha et al. (2015)
15 mins spectrophotometry 100 g−1
Fig pulp and peel TPC and TFC SLE 80% ethanol Vortex mixing for UV/Vis 56.3–285 mg GAE Hoxha and Kongoli
containing 2% 15 min spectrophotometry 100 g−1 (TPC) and (2016)
hydrochloric acid 42.47–
269.54 mg CE g−1
(TFC)
Dried figs TPC and 4 Aqueous Ethanol and Stirring on a magnetic UV/Vis 22.5–288 μg g−1 Feng et al. (2015)
individual two phase dipotassium stirrer spectrophotometry (polyphenols)
polyphenols extraction hydrogen phosphate (TPC) and micellar
solution electrokinetic
chromatography
Dried figs TPC and TFC SLE 70% methanol Maceration UV/Vis 31.8 mg GAE Alamgeer et al. (2017)
spectrophotometry (TPC 100 g−1 (TPC) and
and TFC) and HPLC-UV 538 mg QE 100 g−1
(6 polyphenols) (TFC)
Dried figs TPC and TFC SLE 60% acetone The sample mixture UV/Vis 482–644 mg GAE Bachir Bey and
was shaken at 40 °C spectrophotometry 100 g−1 Louaileche (2015)
for 2 h in a shaking 87.2–126 mg QE
water bath 100 g−1
Fig peels Cyanidin Ultrasound Citric acid acidified The sample was HPLC-DAD-ESI/MS – Backes et al. (2018)
3-rutinoside assisted 100% ethanol ultrasonicated at
extraction 30-35 °C for
approximately 22 min
Dried figs TPC and TFC SLE 80% methanol Samples were UV/Vis 78–458 mg GAE Benmaghnia et al.
(TPC) and 50% macerated under spectrophotometry 100 g−1 (2021)
acetone (TFC) agitation for 24 h 38.8–228 mg CE
100 g−1
Fig pulp and TPC and TFC SLE 100% methanol Maceration at room UV/Vis 0.59–3.82 mg GAE Mahmoudi et al.
peels temperature for 24 h spectrophotometry g−1 (2018)
149–254 mg QE g−1
Fig pulp and 5 phenolic Ionic 1-butyl-3- Ultrasonication for HPLC-DAD 1.99–133 μg 0.1 g−1 Qin et al. (2015)
peels compounds liquid- methylimidazolium 30 min at 30 °C
based hexafluorophosphate
ultrasonic-
assisted
extraction
– Not given
654 B. Moyo and N. T. Tavengwa

trifluoroacetic acid (Debib et al., 2016) and citric acid (Backes et al., 2018; Meziant
et al., 2018).
In order to facilitate the mass transfer of the polyphenols to the solvent when
using SLE, some studies have employed maceration (Alamgeer et al., 2017;
Amessis-Ouchemoukh et al., 2017; Benmaghnia et al., 2021; Mahmoudi et al.,
2018), vortex mixing (Hoxha et al., 2015; Hoxha & Kongoli, 2016), magnetic stir-
ring (Meziant et al., 2018; Pereira et al., 2017) and shaking on a shaker (Bachir Bey
& Louaileche, 2015; Khadhraoui et al., 2019; Nakilcioğlu-Taş & Ötleş, 2021)
(Fig. 28.2). It is noteworthy that the extraction times of these mechanical means of
facilitating extraction varied. Although maceration is a simple method, long extrac-
tion times and low extraction efficiencies are its main disadvantages (Amessis-
Ouchemoukh et al., 2017; Debib et al., 2016). Moreover, the sample needs to be
shaken occasionally to facilitate the mass transfer in this procedure (Njila et al.,
2017). Ultrasound-assisted SLE was the commonly used method to facilitate extrac-
tion in most studies (Ammar et al., 2015; Bachir Bey et al., 2017; Kamiloglu &
Capanoglu, 2015; Viuda-Martos et al., 2015; Wojdyło et al., 2016). Ultrasound
radiation can improve the mass transfer of analytes from the matrix by breaking the
plant material’s cellular walls, subsequently allowing the solvent to penetrate the
matrix (Chen et al., 2007). Furthermore, ultrasonication reduces the matrix particle
size, allowing improved interaction between the solvent and the sample.
Consequently, improving mass transfer results in higher extraction yields in shorter
times (Um et al., 2018). This procedure can also reduce the degradation of thermo-
labile phytochemicals during long extraction processes (Backes et al., 2018).
It is noteworthy that most of the studies reviewed in this paper analyzed total
phenolic content (TPC) and total flavonoid content (TFC). However, these assays do
not give information on the individual polyphenols in the samples. Therefore, to
fully understand the health benefits of figs, the first step is to uncover the phyto-
chemicals that they contain using more sensitive analytical instruments such as liq-
uid chromatography. Moreover, the quantitative analysis of polyphenols from figs
by liquid chromatography is still restricted to a few selected compounds (Alexandre
et al., 2017; Nakilcioğlu-Taş & Ötleş, 2021; Wojdyło et al., 2016).

1.3.2 Modern SLE Based Extraction Methods

Alternative methods to conventional methods, such as high pressure-assisted extrac-

tion, have emerged in the past years to overcome the challenges associated with
traditional techniques for extracting polyphenols from figs (Alexandre et al., 2017).
The main advantage of this technique is that extraction is done at room temperature
hence preventing the thermal degradation of thermo-labile phytochemical com-
pounds. Subcritical assisted extraction (SWE) has also been employed to extract
TPC and TFC. In a study done by Maghsoudlou et al. (2017), SWE was employed
to extract TPC and TFC from fig pulp and skin. As a polar solvent that is inexpen-
sive, available in abundance, and environmentally friendly, water was used in this
study. SWE offers significant advantages over other extraction techniques, including
greater quality extracts, short extraction times, a lower volume of solvents needed,
28 Extraction and Analysis of Polyphenolic Compounds in Ficus carica L. 655

and a low operation cost (Ko et al., 2020). In a study done by Feng et al. (2015), an
aqueous two-phase system was used to recover and partially purify phenolic com-
pounds from fig fruits in a single step. The superiority of this method lies in the mini-
mization of separation stages and high purification of natural products compared to
conventional extraction methods (de Oliveira et al., 2018). Pressurized MAE was
used for extracting bioactive molecules from Ficus carica L. bark (Masota et al.,
2020). However, this method has never been reported for fig fruits. MAE reduces the
thermal gradient created by traditional heating, which augments the degradation risk
of heat-sensitive bioactive compounds (Chaves et al., 2020). In this technique, pres-
sure builds up in the sample matrix, increasing the porosity of the matrix and allow-
ing for improved extraction through solvent penetration (Ameer et al., 2017).
Due to compliance with the principles of green analytical chemistry, green sol-
vents, including ionic liquids (Qin et al., 2015; Xu et al., 2016) and deep eutectic
solvents (Bubalo et al., 2016; Peng et al., 2016; Zannou & Koca, 2020) are increas-
ingly gaining interests in the extraction of phytochemicals. In a study conducted by
Qin et al. (2015), five phenolic acids were extracted using ionic liquid-based
ultrasound-assisted extraction. A small sample mass and volume of the solvent were
used in this study compared to all the studies reviewed in this paper. Extraction was
suspected to be facilitated by multi-interactions between phenolic compounds and
imidazolium cation, including π-π, ionic/charge-charge, and hydrogen bonding.
Remarkably, the application of these non-conventional extraction techniques is still
limited in analyzing polyphenols from fig fruits.

1.4 Methods of Detection of Polyphenols in Fig Fruits

Sample collection, sample extraction, analytical measurement, data processing and

analysis, and biochemical interpretation are common aspects of phytochemical
analysis studies (Aron et al., 2020; Nothias et al., 2020). The analytical workhorses
of phytochemical analysis studies are mass spectrometry (MS) and nuclear mag-
netic resonance (NMR) spectroscopy (Stefanucci et al., 2020; Amaar et al., 2015;
Nhoek et al., 2021). The sophisticated analytical technique of liquid chromatography-
mass spectrometry (LC-MS) identifies and quantifies a broad spectrum of metabo-
lites. Its benefits include sensitivity and a quick and easy procedure for large-scale
sample analysis, making it the instrument of choice for untargeted metabolomics of
plant-derived natural compounds (Salem et al., 2020). Despite these developments
in analytical instrumentation, spectrophotometric methods are still widely used to
date for the determination of polyphenols from figs (Ammar et al., 2015; Bachir
Bey & Louaileche, 2015; Debib et al., 2016; Feng et al., 2015; Hoxha et al., 2015;
Kamiglou et al., 2015; Viuda-Martos et al., 2015; Harzallah et al., 2016; Hoxha &
Kongoli, 2016; Alamgeer et al., 2017; Amessis-Ouchemoukh et al., 2017; Alexandre
et al., 2017; Ersoy et al., 2017; Konak et al., 2017; Maghsoudlou et al., 2017; Pereira
et al., 2017; Mahmoudi et al., 2018; Meziant et al., 2018; Mopuri et al., 2018;
Khadhraoui et al., 2019; Benmaghnia et al., 2021; Nakilcioğlu-Taş & Ötleş, 2021).
The Folin-Ciocalteu method, a colometric assay, is the commonly used method for
656 B. Moyo and N. T. Tavengwa

TPC, whereas the content of flavonoids is usually determined as the total flavonoid
content, spectrophotometrically also (Kamiloglu & Capanoglu, 2015; Khadhraoui
et al., 2019; Maghsoudlou et al., 2017). It is noteworthy that, despite its widespread
usage, the Folin-Ciocalteu method is susceptible to interferences contained in plant
samples, including sugars, ascorbic acid, proteins, and non-phenolic organic com-
pounds that react with the Folin-Ciocalteu reagent (Arvaniti et al., 2019). A limited
number of studies have used liquid chromatography to survey the polyphenols pres-
ent in figs (Ammar et al., 2015). In a study done by Amaar et al. (2015), 91 phenolic
compounds were identified using the metabolic profiling approach. It is noteworthy
that this is the only study where the metabolite profile of polyphenols was investi-
gated in-depth of all the reviewed studies in this chapter.

2 Challenges and Future Trends

Organic solvents are still extensively used to extract various polyphenols in the SLE
procedure, and these solvents are associated with various challenges, including
toxic residues, chemical alteration of extracts, and generation of toxic waste
(Amessis-Ouchemoukh et al., 2017; Mopuri et al., 2018). Moreover, traditional
extraction methods, including SLE, have low extraction efficiencies. As a result,
innovative techniques that prevent harmful solvents while enhancing efficiency and
sustainability are required (Vernès et al., 2020). Significant efforts have been made
in this direction, and to date, a significant number of innovative extraction tech-
niques using green and environmentally friendly solvents, such as deep eutectic
solvents (Bubalo et al., 2016; Peng et al., 2016; Zannou & Koca, 2020) and supra-
molecular solvents (Cai et al., 2022; Torres-Valenzuela et al., 2020) have been
made. Moreover, modern extraction techniques that use water as an extraction sol-
vent, such as PHWE, have been introduced in the past years (Cvetanović et al.,
2018; Lachos-Perez et al., 2020). Furthermore, miniaturized extraction techniques
such as dispersive liquid-liquid micro-extraction (DLLME) (Diuzheva et al., 2018;
Fusari et al., 2019; Ramirez et al., 2021), hollow fiber liquid micro-extraction
(Mlunguza et al., 2020; Moyo & Tavengwa, 2021b) were developed in the past
decade, and they consume less volumes of organic solvents. Additionally, DLLME
has short extraction times. Therefore, future research on the extraction of phyto-
chemicals from figs should consider employing these modern extraction techniques
and green solvents.
The high concentration of free sugars and other matrices can challenge analyzing
phenolic compounds from figs using the SLE technique (Arvaniti et al., 2019).
Therefore, compounds other than the analytes of interest and other interfering sub-
stances may need to be removed via an additional clean-up technique such as solid-
phase extraction. Moreover, selective sorbents such as molecularly imprinted
polymers may reduce the matrix interferences (Selvamuthukumaran & Shi, 2017).
Currently, there is scanty information on the specific phytochemicals found in fig
fruits (Alamgeer et al., 2017; Ammar et al., 2015; Feng et al., 2015; Wojdyło et al.,
2016). As a result, more profiling studies should be conducted, and this may uncover
new metabolites that may be potent besides the widely studied polyphenolic classes.
28 Extraction and Analysis of Polyphenolic Compounds in Ficus carica L. 657

3 Conclusion

The most studied phytochemical compounds present in fresh and dried figs are phe-
nolic acids and flavonoids. SLE is still the widely employed extraction technique for
polyphenols from figs, despite its disadvantages. Future research should consider
modern extraction techniques and green solvents as they offer various advantages
over conventional methods. Colorimetric assays are still widely used to determine
total polyphenolic compounds in these samples. Despite being well-studied, the
chemical make-up of the fig tree needs to be further investigated; mainly, the meta-
bolic profiling approach can be used to decipher diverse phytochemicals present in
these fruits. Supplementing traditional assays with more specific and selective ana-
lytical instrumentation might help understand the health benefits of these fruits and
potentially potent drugs that might be discovered via this route. Concentrations of
phenolic compounds vary, and this might be due to the methods of extraction and
detection used, season, level of maturity, and fig variety.

Conflict of Interest The authors declare no conflict of interest.

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Part III
Fig (Ficus carica): Technology, Processing,
and Applications
Chapter 29
Fig (Ficus carica) Drying Technologies

Olfa Rebai, Oumayma Ghaffari, and Sami Fattouch

Abbreviations

1-MCP 1-Methylcyclopropene
AD Air-convection drying
MAP Modified atmosphere packaging
MW Microwave
MWAD Microwave-air convection
MWVD Microwave vacuum drying
T.S.S Total soluble solid
TA Titratable acidity
VD Vacuum-drying

1 Introduction

Fig (Ficus carica L.) belongs to the order of Urticales and the family of Moraceae
with over 1400 species. One of the first worldwide cultivated trees for its edible
fruit. It is supposed to originate from Western Asia, spread to the Mediterranean by
humans, and grow in many parts of the world with moderate climates (Baraket et al.,
2009). Besides its primary metabolites content, several phytochemical studies on
F. carica revealed the presence of a panoply of bioactive phenolics, phytosterols,
organic acids, anthocyanins, triterpenoids, coumarins, and volatile compounds such

O. Rebai (*) · O. Ghaffari · S. Fattouch

EcoChem Laboratory, Department of Biological and Chemical Engineering,
National Institute of Applied Sciences and Technology (INSAT),
University of Carthage (UCAR), Tunis, Tunisia
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 665
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_29
666 O. Rebai et al.

as aliphatic alcohols, and few other classes of secondary metabolites (Oliveira et al.,
2009). Figs contain essential amino acids and are cholesterol-free. They are an
excellent source of minerals, vitamins, carbohydrates, and dietary fiber.
Thanks to their wide variety of chemical constituents, Ficus carica L. fruits are
used in traditional medicine to remedy many health problems and biological activi-
ties. F. carica has been traditionally used for its medicinal benefits as a metabolic,
cardiovascular, respiratory, antispasmodic, and anti-inflammatory remedy (Duke et
al., 2002). Leaves and roots of F. carica are used in different disorders such as gas-
trointestinal (colic, indigestion, loss of appetite, and diarrhea), respiratory (sore
throats, cough, and bronchial problems), inflammatory, and cardiovascular disor-
ders. Figs have a laxative effect due to their content of many antioxidants. Moreover,
they are a good source of flavonoids and other polyphenols. The phenolic antioxi-
dant qualitative and quantitative composition of dried fig fruits was determined and
analyzed by several studies and investigators. The F. carica juice mixed with honey
is commonly used for hemorrhage. In Indian medicine, fig fruits are a mild laxative,
expectorant, and diuretic (Solomon et al., 2006).
F. carica fruits could be used fresh, dried, canned, or as jam (Tous & Fergueson,
1996). Recurrent consumer anxiety is rising about using preservatives and chemical
additives for food storage. Thus, many efforts have been made to find safe preserva-
tion methods during the last decades. Fruits and vegetables contain a large quantity of
water, due to which they rapidly deteriorate. Drying is among the most used processes
to remove the food’s moisture, so bacteria, yeast, and mold cannot grow and spoil the
food. The nutritional value of food is only minimally affected by drying (Ahmed et
al., 2013). During water evaporation, the fruit decreases in weight and shrink; how-
ever, its beneficial substances do not disappear and are “preserved”. Generally, dried
foods are tasty, nutritious, lightweight, and easy to prepare, store and use. A fresh fig
contains total sugar (mainly glucose and fructose), but it rises to about 50% when
dried. It is commercialized in the market as sweet due to its high level of sugars and
provides an excellent dietary substitute (Veberic et al., 2008). Paradoxically, fig pre-
served forms, especially dried fruits, are used as supplement food for diabetics in
some eastern countries. Moreover, dried flesh has been known as a source of medi-
cine or a part of medicine in many Mediterranean countries, i.e., in Turkey, Egypt,
Tunisia, Algeria, Morocco, Spain, Greece, and Italy, since the old time.

2 Criteria to Consider for Dried Fruits

Food drying is one of the oldest methods of preserving food for later use. It could
either be an alternative to canning, freezing, or complimenting these methods.
Drying removes excess (free) water, compromising the sensory and nutritive values.
By definition, food dehydration removes free water from food, generally by circu-
lating hot air through it, which prohibits the growth of spoiling microbials, particu-
larly bacteria. Drying also slows down the action of enzymes (naturally occurring
substances that cause food to ripen) but does not inactivate them. When water is
removed from a material, its structure changes and becomes harder and less likely
29 Fig (Ficus carica) Drying Technologies 667

to spoil. When dried, the food becomes smaller and lighter in weight. When needed,
in some cases, the water is added back to have food ready for use, allowing the food
to return to its original shape and texture.
Drying empirically uses heat to remove water from a raw or processed material.
This technological method can be done in several ways, including solar drying or
using an oven, a microwave, or a sophisticated dehydrator. The sun, the wind, and a
smoky fire removed water from fruits, meats, grains, and herbs. The energy input is
less than needed to freeze, and the storage space is minimal compared with that
needed for canning jars and frozen containers. Foods can be dried using warm tem-
peratures, low humidity, and air current. In drying, warm temperatures cause the
moisture to evaporate. The higher the temperature gradient between the food and
the ambient air is, the increased speed in the evaporation of food water. However,
when considering hysteresis phenomena, an excess temperature gradient could
harm the food texture. Generally, a food-specific temperature regime should be set
and optimized. Commonly, heating at medium (50–55 °C) or relatively high (60 °C)
temperatures is used. In addition, lower air humidity allows moisture to move
quickly from the food to the air. The heat and mass (water) transfer depend on the
velocity of the circulating air. The air current speeds up drying by moving the sur-
rounding moist air away from the food.

3 Fig Drying Technologies

Fig fruit (Ficus carica L.) could be dried either by traditional methods (sun or solar
drying) or in conventional hot-air dryers (Babalis et al., 2006). Drying might be
classified into two groups: natural drying and artificial drying. Mechanical dehydra-
tion or artificial dehydration can be further classified into atmospheric and sub-
atmospheric types based on the conditions employed in the drying process. Artificial
drying is done with the help of mechanical or electrical equipment. Based on the
model of the drying process, drying at atmospheric pressure conditions could be
further divided into batch and continuous types. Mechanical drying includes the
methods of drying by (1) heated air, (2) direct contact with a heated surface, e.g.,
drum drying, and (3) application of energy from a radiating microwave or dielectric
source. According to Ahrens et al. (2000), artificial methods could efficiently
remove a large amount of moisture. In addition, controlling various factors such as
temperature, drying air flux, and drying time is also possible when artificial meth-
ods are used (Maisnam et al., 2017).

3.1 Natural Sun-Drying

Drying is one of the oldest preservation processes available to humankind that we

could track since prehistoric times. People have sun-dried fruit and vegetables for
thousands of years to preserve for leaner times. The relatively high sugar and acid
668 O. Rebai et al.

content of fruits was enough to make them safe to dry in the sun for a few days. New
technologies brought changed techniques, but the increasing demand for healthy,
low-cost natural foods and the need for sustainable income, are bringing solar drying
to the fore as a valuable alternative for surplus products. Sun-drying works best in
hot, dry climates, but it can also be successful in more humid zones. In today’s food
market, dried food plays an essential role in the food supply chain. As for fruits and
vegetables, it can be estimated that they constitute about 1% of the total drying in the
food industry, by large being the grains the most important (Ahmed et al., 2013).
Sun driers are classed according to the systems of heating the product. They are
not designed based on heat and mass transfer calculations or principles since they
remain traditional and artisanal.
(1) Natural drier is the traditional common means which uses the direct sun action
on the materiel itself and the ambient air to which the product is exposed. In
these driers, the raw products are spread out in an enclosed space where they are
protected against dust contamination and subjected to limited airflow control
(just using some physical obstacles to break strong air current, if needed).
(2) Direct driers: The material to be dried in these units is placed in an enclosure
with a transparent cover. Heat is generated by sun radiation of the product itself
and the internal surfaces of the enclosure. The heat causes the moisture to evap-
orate from the product. In addition, it expands the air in the enclosure, creating
air circulation, which facilitates mass and energy (heat) transfers.
(3) Indirect driers: The sun radiation heats the air exclusively in these driers. The
hot air is then led by thermocirculation or by forced circulation into a chamber
or cabinet holding the products to be dried. Different instruments of this type
are available in the market; some are associated with a fan to intensify and con-
trol air circulation.
(4) Mixed driers: are indirect driers where the hot air from the exterior solar heater
is combined with direct solar heating of the products through a transparent roof
of the drying cabinet (FAO, 1985).
The drying process of food products is to leave them outside in the sun and the
windy position and allow them to dry within 7–10 days, depending on the air cur-
rent, temperature, and humidity, for good dehydration. As far as drying temperature
is concerned, the most important thing is to dry them in sunny and dry weather,
particularly in a hot summer. Drying in the sun is very economical. Operators only
have to spread, as much as possible, the produce on a suitable surface and let it dry
in the sun. The nutritional benefits of sun drying, or solar drying, are that it takes just
enough moisture out of fruits to prevent them from spoiling while being stored.
Sun-drying retains large amounts of vitamins, fiber, and minerals. For people with
diabetes, dried fruit prepared without adding sugar (water activity, aw, depressor) is
a healthy choice instead of desserts (Dhaouadi et al., 2014).
Sun-dried fig (Fig. 29.1) can be eaten from starter to dessert. It can be eaten plain
or stuffed with almonds or walnuts. It also goes perfectly with the flavors of
Mediterranean cuisine, especially in North African “couscous” and “tajines”. Cut
into small pieces; it is a wonderful addition to salads. In addition, it improves the
bargaining position of farmers. Sometimes farmers sell figs at very low prices
29 Fig (Ficus carica) Drying Technologies 669

Fig. 29.1 From left to right, whole F. carica fruit, the fruit cut open, and sun-dried fruits (Sukowati
et al., 2019)

during the harvest season because they cannot store or preserve their surplus prod-
ucts, thus encouraging people to establish their sun-dried figs.
However, disadvantages associated with these traditional sun dryers are well
known and should be highlighted here, that it is an uncontrolled method. Mainly, fig
fruits did not dry uniformly, thus leading to heterogeneously stabilized/unstabilized
products during storage. The drying is slow and sometimes incomplete under unfa-
vorable climatological conditions. When the sun is not available, this method is not
possible. Moreover, this process requires many unskilled laborers, takes a long time,
and requires a large area. Constantly, sun-dried figs are subjected to noxious effects
of dust, dirt, and insect infestation. Regular and incessant supervision of the dryer is
necessary as the product must be gathered and moved undercover in the event of
rain, and predators must be chased away. Indeed, possible losses of crops by birds,
hens, or other animals are inevitable.

3.2 Artificial Drying

The method in which excess water of figs is removed artificially offers better control
than natural drying, thus, resulting in greater product uniformity and quality.
However, the initial investment in equipment and expenditures on energy inputs are
high and may not always be justified. Therefore, different types of artificial drying
are used: (1) mechanical drying, (2) infrared or (3) dielectric drying, and (4) chemi-
cal drying.

3.2.1 Solar Drying

Using a solar drying system for fruits is becoming an attractive alternative food
preservation technique over conventional ones. Solar drying also uses the sun as the
heat source. Therefore, solar drying remains a viable drying and food preservation
technology alternative with the potential to reduce post-harvest losses, which can be
as high as 25–30% in food grains and 30–50% in fruits and vegetables, as reported
670 O. Rebai et al.

Fig. 29.2 Experimental setup of a cabinet type drier (Phadke et al., 2015)

by Patel et al. (2013). This is due to many advantages of the solar drying system
compared to others: low cost, high quality of dried products, and environmentally
friendly (Heredia et al., 2007). Here, instruments are designed based on heat and
mass transfer calculations and optimization (Fig. 29.2). Generally, three types of
artificial solar driers are used, (1) the absorption or hot-box type driers in which the
product is directly heated by the sun, (2) the indirect or convection driers in which
the product is exposed to warm air which is heated using a solar absorber or heat
exchanger and (3) drier, which is a combination of the first and the second types. In
solar drying, the energy source is the sun. Therefore, it is a very cheap drying
method, but on the other hand, it has many drawbacks because the food is exposed
to contamination sources (insects, birds, and other animals) and is also strongly
susceptible to weather conditions. Indirect solar drying, the product is exposed to
solar radiation, whereas, in indirect solar drying, the sun’s energy is harnessed by
collectors used to heat the air that will contact the fig fruits to dry (Patel et al., 2009).

3.2.2 Heat Pump Dryer

Heat pump fruit drying is used to dry all kinds of food products in hygienic ambient
at economic conditions with low energy consumption. Due to the increasing prices
of fossils and electricity and the emission of CO2 in conventional drying methods,
green energy-saving and other heat recovery methods for processing and drying
food have become very important. Heat pump technology has been successfully
used for drying agricultural products and other domestic dehumidification/heating
applications. It has excellent prospects and revolutionary ability. However, heat
pump drying (HPD) of fruits and vegetables has been largely unexploited in South
29 Fig (Ficus carica) Drying Technologies 671

Africa and, by extension, in the sub-Saharan African region (Fayose & Huan, 2016).
Fig drying using a heat pump reduces energy cost with relatively low air tempera-
ture and relative humidity. This process could be achieved by condensing moisture
from the drying air and then reheating, and product quality can be improved in
terms of color and smell (Soponronnarit et al., 2007).
HPD technique is more effective in drying material with more free moisture in
some fruits. The application of heat pump drying contributes positively to dried fruit
and vegetables’ quality attributes, including improved microbial safety, better color,
vitamin C retention, enhanced volatile compounds, aroma and flavor substances, rehy-
dration, and texture. In addition, compared to sun drying, product quality is improved
in preserving the product’s natural sensorial properties (Soponronnarit et al., 2007).

3.2.3 Pilot Cabinet Dryer

Cabinet dryers are the most popular devices for fruit drying. However, one of the
drawbacks of this dryer can be nonuniformity in the desired end product moisture
content (Darabi et al., 2015). The cabinet dryer tested is of the type pioneered by the
Brace Research Institute in Canada (Anon, 1965). It is essentially a rectangular
container insulated at its base and covered with a glass or transparent plastic roof.
Holes drilled through the base permit fresh air to enter the cabinet. Outlet holes are
located on the upper parts of the cabinet side and rear panels. In this case, the cabi-
net was constructed of thick plywood panels.
Mujić et al. (2012) used a pilot plant cabinet dryer to compare the aroma profiles
of fresh figs and figs frozen in liquid nitrogen and non-treated and pre-treated dried
figs of the Croatian cultivar. This study offers an insight into volatiles preserved dur-
ing drying, and results have shown that pretreated dried figs with sulfur dioxide were
capable of preserving many aromatic compounds that are important for consumers.

3.2.4 Microwave Drying

Called also drying by radiation. Generally, long drying time and high temperature
are the factors responsible for the loss of heat-sensitive components of the food.
However, microwave drying has been considered excellent in retaining nutritional
and sensory values besides saving energy and time in recent years. Hence, micro-
wave drying is used by several modern food industries worldwide. Microwaves
(MWs) are electromagnetic (EM) waves synchronized with perpendicular oscilla-
tions of electric and magnetic fields in a frequency between 300 MHz and 300 GHz,
ranging from 1 m to 1 mm. Microwave heating mechanisms are based on the oscil-
lation of ions and polar molecules mainly water when a material is exposed to the
EM waves which cause the internal friction and conversion of kinetic energy into
heat (Laguerre et al. 2018).
During microwave drying, the quality preservation of the product is of utmost
importance (Khraisheh et al., 2004). Product quality includes three principal areas:
nutritional value, acceptability, and safety. The reasons for uneven heating with MWs
are due to several factors: (i) large product size; (ii) resonance phenomena; (iii)
672 O. Rebai et al.

heterogeneous material composition, and (iv) shape of the product (Radojčin et al.,
2021). Exposure of food to infra-red led to charge build-up in the electronic state and
the vibrational and rotational state at the atomic and molecular levels. This causes the
food to heat up without any changes in the air temperature surrounding the food.
Sharifian et al. (2013) investigated the qualitative indices for the pulsed microwave
dried figs (Ficus carica L.) through image processing techniques. Kinetic parameters
for the color change were determined using the total color change parameter, chroma,
hue angle, and browning index. Also, in the same study, Sharifian et al. (2013) dem-
onstrated that the MW power intensity and pulsing ratio significantly influenced the
color change of fig fruit during drying. Increasing MW power intensity can provide a
desirable contribution to the light yellowish color of dried figs. Nevertheless, from the
growing total color changes, the authors concluded that 2.5 W/g is the optimum MW
power intensity level concerning fig color criteria (Sharifian et al., 2013). However,
depending on the type of material, MWs, in some cases, cannot fully complete drying
and are usually combined with hot air drying or vacuuming.
Sharifian et al. (2012) investigated the effects of microwave power intensity and
pulsing ratio on the drying behavior of fig fruits in a laboratory-scale microwave
dryer and has shown that the drying time of products increased by approximately
200% under the pulsing ratio of 1.5–4. In contrast, drying time decreased by approx-
imately 500% under microwave power intensity of 0.5–2.5 W/g; microwave power
intensity resulted in the raised sample temperature leading to better moisture
removal. Sample temperature decreased with the pulsing ratio while increased with
MW power intensity. The warming-up period was short, and subsequent drying
only occurred during the continuous temperature.

3.2.5 Microwave Hot Air Drying

Microwave heating is used in many processes such as baking, pre-cooking, and dry-
ing industry. This method associates the MW process and hot air circulation as a
mass and heat transfer vector. The microwave energy is absorbed by food materials
and converted into heat due to the ionic interaction and the dipolar rotation. Salt and
water are common molecules in most foods. Therefore, salt molecules are vibrated
due to ionic interaction. The application of microwave heating in conjunction with
hot air drying has led to a significant reduction in the drying time of fig fruits (Abul-
Fadl et al., 2015).

3.2.6 Microwave Vacuum Drying

Giri and Prasad (2007) reported that food products’ quality could be better main-
tained by vacuum drying. However, vacuum drying requires high investment and
operating costs and thus is only suitable for high-value products such as medicinal
herbs. In addition, vacuum drying has a poor mass transfer rate and leads to an
extended drying time (Wang et al., 2018). Correspondingly, Xu et al. (2006) reported
that vacuum drying has high operating costs due to the need to maintain a vacuum
over long drying periods.
29 Fig (Ficus carica) Drying Technologies 673

3.2.7 Heat-Drying Ovens

An oven is a direct-fired chamber or cabinet holding the products to be dried that

distributes heat effectively. Ovens are widely used in the food industry to preserve
perishable commodities, especially foodstuff and heating and drying. The oven con-
sists of an insulator box that serves as the housing for which the food is placed for
baking, a heating element, and a thermostat that regulates the heat generation rate and
regulates the fuel or current flow to the heating element. Modern ovens are fueled by
gas, electricity, or both and are commonly used for cooking, frying, baking, and roast-
ing. However, the daily need, high market demands, and problems often associated
with the high cost of procuring the gas or kerosene type of oven for cooking, drying
and baking prompt us to develop an electric-powered oven for baking bread cake etc.

3.2.8 Electric Drying

Many studies have shown that nonthermal food processing techniques can nega-
tively affect food quality. The first commercial electric drier was developed in 2008,
which has been improved to exceed hundreds of kg of product to dry in one batch
(Martínez-García et al., 2013a, b). No fuel is used, no CO2 emission, thus making
this process eco-friend. The electric drier is a heating chamber meant for evaporat-
ing food’s water by converting electrical energy to heat or thermal energy. These
novel pretreatment and treatment techniques have been developed further to improve
product quality (Deng et al., 2017). With no waste heat loss, the circulating air vol-
ume can be reduced by 20 to 40% compared to other conventional driers. When
miniaturized, an electric dryer can be portable, relatively cheap, readily available,
and easy to install, operate and maintain (Adegbola et al., 2012). Nowadays, these
instruments are more and more electronically controlled.
Electric dryers work on the principle of convection or infrared radiation. The hot
air convection makes the food moisture gradually released. Even though this dryer
has an affordable cost, it may induce fruit deformation, change its taste, and lose
some vitamins. Infrared-based electric dryers use radiations that penetrate deeply
into fruits, so the drying occurs at sufficiently low temperatures, not exceeding
60 °C. The fruit’s practically preserved taste, color, shape, and vitamins (preserved
up to 90%) make this electric drying process ideal.

3.2.9 Tray Dryer

A tray dryer is convectional drying equipment with enclosed insulated chambers

and trays in a trolley placed on top of each other. The application of tray dryer is
widely used in agricultural drying because of its simple design and capability to dry
products at high volume. However, the greatest drawback of the tray dryer is uneven
drying because of poor airflow distribution in the drying chamber (Misha et al.,
2013). Gwala and Padmavati (2016) compared the drying kinetics of an electric tray
drying, a solar cabinet drying, and an open sun drying. The tray dryer was superior
674 O. Rebai et al.

in heat transfer but weaker in mass transfer, explaining the little difference observed
in drying time compared to solar and sun drying. The evidence was suggested by
drying models, drying constants, and the effective moisture diffusivity model
(Gwala & Padmavati, 2016).

3.2.10 Spray Drying Technology

Spray drying equipment and techniques developed over several decades, from the
1870s through the early 1900s. Spray drying comes of age during World War II, with
the sudden need to reduce the transport weight of foods and other materials (Sonone
et al., 2016). This method transforms feed from a fluid state into dried particulate
form by spraying the feed into a hot drying medium (Andreou et al., 2021). It is a
continuous particle processing drying operation. The feed can be a solution, suspen-
sion, dispersion, or emulsion. The dried product could be in the form of powders,
granules, or agglomerates, depending upon the physical and chemical properties of
the feed, the dryer design, and the final powder properties desired (Killeen, 2000).
The spray drying process has advantages that could be designed to virtually any
capacity required. Feed rates range from a few pounds per hour to over 100 tons per
hour. Operation is continuous and adaptable to fully automatic control (Gharsallaoui
et al., 2007). It can be used with both heat-resistant and heat-sensitive products
(Chegini & Ghobadian, 2007). Nearly spherical particles can be produced. Some
limitations include limited versatility in producing particles or structures with com-
plex morphologies and rapid drug release rates often exhibiting a burst effect.
Due to existing challenges in the field of syrup or powder production and the
presence of strong demand for turning waste into high-value products. Kalantari et
al. (2018) examined the production of fig extract from waste figs under mild extrac-
tion conditions and attempted to optimize fig syrup’s extraction and drying condi-
tions. Fig extract was used to feed the spray dryer to prepare a powder with suitable
physicochemical properties to produce seedless fig paste or drink. Furthermore, the
effects of drying aids and spray drying operating conditions were investigated on the
quality of the obtained powder. Results show that the optimal extraction conditions
for producing fig extract are suggested to be 5 °C at 72 h. Fig extract powder was
produced from the natural fig extract using a pilot-scale two-fluid nozzle spray dryer.
Furthermore, increasing the inlet air temperature led to increased porosity, dis-
persibility, and particle size distribution. In addition, an increase in the airflow rate
increased the moisture content, dispersibility, and porosity of the powders. The best
powder was produced at a temperature of 170 °C using an airflow rate of 400 m3/h,
conditions that can be a good strategy for converting waste figs to high-value prod-
ucts capable of being used to formulate healthy food products Kalantari et al., 2018.
The quality characteristics of spray-dried powders from unripe fig extract were
investigated by Chae and Hong (2016). The spray-dried powder showed the highest
protease activity with (0.84 unit/g), moisture content, and L value of the spray-dried
powder than those the freeze-dried powder. Furthermore, in the same study, in vitro
digestion experience, spray-dried powders of the unripe fig showed a protease sur-
vival until approximately 25% (Chae & Hong, 2016).
29 Fig (Ficus carica) Drying Technologies 675

4 Advantages and Disadvantages of Fig Drying

Drying is one of the oldest unit operations and has recently become widespread. It
has been used in different industries to gain different utilities. The methods of dry-
ing are diversified according to the purpose of the process. There are more than 200
types of dryers. For every dryer, the process conditions, such as drying chamber
temperature, pressure, air velocity (if the carrier gas is air), relative humidity, and
product retention time, have to be determined according to feed, product, purpose,
different drying methods used to dry and retain the quality of the dried fig fruit.
Therefore, there is a greater need to select an appropriate drying method for good
quality figs considering prevailing conditions. The objective is the different drying
methods to dry the fig fruit to increase its shelf life and select appropriate drying
methods for figs of good quality.

4.1 Impact of Fig Drying on Its Technological Characteristics

Drying food is a process by which water is removed from the food by vaporization
or sublimation, thus reducing the water available for chemical, enzymatic or micro-
bial degradation reactions. The acceptability (visual appeal, taste, aroma, flavor, and
texture), structural property, and nutritional value of fruits and vegetables are also
highly affected (Zhang et al., 2005; Zhang & Xu, 2003). Thus, the main objective of
any drying process is to produce a dried product of desired quality at minimum cost
and maximum throughput and to optimize these factors consistently (Chua et al.,
2001). Technological treatment changes the structure of the raw fruit by modifying
enzymatic reactions occurring in the tissue and affecting the heat and mass exchange
conditions in the plant material (Janowicz et al., 2008). Each drying technique
depends on various factors, such as the required type of product, size, level of ripe-
ness, structure, color, aroma, chemical composition, nutritional composition,
expected final quality, availability of a dryer, and costs. These changes and influenc-
ing factors are physical quality, chemical quality, nutrition, and sensory quality. The
main objectives of drying include preserving food and increasing its shelf life by
reducing the water content and water activity; avoiding the need for the use of
refrigeration systems for transport and storage (expensive); reducing space require-
ments for storage and transport; diversifying the supply of foods with different fla-
vors and textures, thus offering the consumers a great choice when buying foods
(Guiné, 2018).
The drying conditions used, mainly heat and humidity can significantly influence
fig fruit’s technological and functional characteristics. Effect of drying on physico-
chemical, functional, and morphological characteristics of the fig. Generally, the
process of drying in spouted beds has been presented in the literature as an attractive
alternative for drying pastes and suspensions, producing a high-quality powder at
and low cost. It is widely used in the dehydration of heat-sensitive materials to pre-
serve bioactive compounds and other structures of interest, offering advantages over
other methods of drying, especially short drying times (Bezerra et al., 2013).
676 O. Rebai et al.

The intensity of changes in quality parameters during drying depends on the

nature of the material, pretreatment, used process, and their parameters. These
changes include color, shape, volume, density, texture, flavor, nutrients, water activ-
ity, microbiology, rehydration ability, etc. Ong & Law, 2010. Convective drying
kinetics is affected by many factors, such as fig size and composition and thermo-
dynamics properties of drying air such as temperature, humidity, and velocity. With
drying, fig mass and volume are reduced, and packaging and handling costs. It also
reduces the microbiological activity and chemical changes during the storage of the
final dehydrated product (Martínez-García et al., 2013a, b).
Nagaraja et al. (2016) evaluated different drying methods for pre-treated fig
fruits. Fig fruit was dried under different drying methods (sun drying, solar cabinet
dryer, microwave oven, and cabinet tray dryer) to elicit their effect on drying rate,
drying time, biochemical properties, and sensory attributes. Microwave oven drying
showed the highest drying rate and took less time to reach the required final mois-
ture content. A higher drying rate was observed in the pre-treated samples than in
the untreated samples under all drying methods. The drying time for cut-fig fruits
was nearly half of that for whole fig fruits. The sulphited samples showed higher
biochemical properties irrespective of drying methods. The samples dried in a
microwave oven, solar cabinet, and sun-drying had higher values of biochemical
properties than the cabinet tray dried samples. Acceptable sensory quality was
obtained in products dried with a microwave oven, solar cabinet, and sun-drying
(Nagaraja et al., 2016). Babalis studied the influence of drying air temperature and
air velocity, and Babalis and Belessiotis (2004) demonstrated a strong influence of
air temperature and velocity at the early drying stages. The relative insensitivity of
the drying process at the later stages. Higher air temperatures affect significantly the
drying rate at the first stages of drying, but their relative effect diminishes after a
period of about 10–15 h. Then, a range of 1–2 m/s for the drying air velocity has to
be preferably established inside an industrial dryer for agricultural products, giving
the best cost/efficiency performance for the drying of the figs. These investigations
revealed that the drying kinetics is most significantly affected by temperature, with
the air flow velocity having a limited influence on the drying process. Correlations
expressing the drying constants and effective moisture diffusivity dependence on
the drying-air parameters are also reported (Babalis & Belessiotis, 2004).
Another trend in drying technology is Osmotic dehydration; OD is the process of
water removal by immersion of water-containing cellular solid in a concentrated
aqueous solution (Ponting, 1973). The fundamental purpose of food dehydration is
to lower the water content to minimize rates of chemical reactions and facilitate
distribution and storage. In osmotic dehydration, foods are immersed or soaked in a
saline or sugar solution. This results in three counter mass transfer phenomena
(Ponting, 1973).
Martínez-García et al. (2013a, b) investigated drying parameters on figs and their
functional properties. They demonstrated that the drying process increased the total
phenolic content, degraded the anthocyanin content of figs, and enhanced the dried
fig's antioxidant activity. Furthermore, dehydrating ground figs was faster and main-
tained their functional properties better than half-cut figs.
Physical properties of food products are responsible for perceived quality; the
color of food is measured directly and non-destructively using apparatus like
29 Fig (Ficus carica) Drying Technologies 677

colorimeters. They use several color spaces for the expression of color values. The
usually used color space is CIELAB (CIE L*a*b*, color system), and this method
helped monitor color changes during the drying of fig. Texture properties include
structural and mechanical characteristics that significantly influence dried products'
quality. Abul-Fadl et al. (2015) investigated the effect of different drying methods
on the quality criteria of dried Fig fruits. In this study, the relation between moisture
content and elapsed time to reach the required moisture content (drying curves) of
dried tested fig fruits for air convection (AD) were compared with combined micro-
wave-air convection (MWAD) drying methods. A combined microwave-hot air dry-
ing (MWHA) technique may be a better alternative. This method provides a higher
drying rate and a better product quality than microwave drying or hot air drying
(Chandrasekaran et al., 2013).
The final drying time of fruits fig using air convection drying AD and vacuum
drying VD methods compared to the combined microwave-air convection MWAD
and microwave- vacuum drying MWVD methods indicated that the much longer
drying time was obtained by using the vacuum drying method for fig. These results
may be due to a slight mass transfer rate in the vacuum drying method leading to a
higher drying time in tested samples than in other drying methods used (Abul-Fadl
et al., 2015).
The chemical composition of fresh and dried fig was also reviewed by Arvanitia
et al. (2019), presents the main phytochemical compounds found in fresh and dried
figs of different varieties, describes the analytical methods used for their determina-
tion, and discuss the antioxidant capacity and the potential effects of figs in
human health.
Abul-Fadl et al. (2015) have investigated results concerning the effect of differ-
ent drying methods on the chemical composition of dried figs produced. No signifi-
cant differences were observed between fig samples dried by AD and VD compared
with the samples dried by the combined MWAD and MWVD in moisture, ash, and
crude fiber contents. On the other side, significant differences were found between
the dried fig by using the AD method and the samples dried by MWAD, and also
with the tested samples dried by using VD and MWVD of protein, lipid, and total
sugars significantly reducing sugars. At the same time, the tested samples dried by
using AD have the lowest content of the previous components of protein, lipid, total
sugars, and reducing sugars.

4.2 Effects on the Chemical Composition and

Nutraceutical Content

It is already known that drying fruit and vegetables using high temperature and for
a long drying time by conventional heating damages the quality of the final dried
products (Viswanathan et al., 2003). Almost all dried fruits provide essential nutri-
ents and an array of health protective bioactive ingredients that help to reduce the
risk of illness by preventing chronic diseases. Natural products have the potential to
be used as therapeutic drugs for humans and livestock species. Along with their
678 O. Rebai et al.

analogs, such compounds can also act as intermediates to produce valuable drugs
(Makkar et al., 2009).
Fig (Ficus carica L.) is one of the most important agricultural products of the
tropic and subtropics areas. In the Middle East and the Mediterranean region, the fig
has been included in the diet since the ancient years, and it is considered the symbol
of longevity. In the northern Mediterranean region, fig trees produce one or two
crops per year, depending on the cultivar.
In general, most of the monitoring studies have demonstrated that the phyto-
chemical composition is often affected by the fig variety, but it is also strongly
dependent on other factors such as the color, the part of the fruit, the level of matu-
rity, and also the drying process that has been used (Harzallah et al., 2016).
The fig is a healthy fruit with numerous medicinal properties that may reduce the
risk of cancer and cardiovascular diseases. Fig fruit is consumed fresh, dried, pre-
served, canned, and candied. It is used for alcohol and wine production in the
Mediterranean region and Europe for fig-coffee preparation. Fresh and dried figs are
especially rich in fiber, trace minerals, antioxidant polyphenols, proteins, sugars,
organic acids, and volatile compounds that provide a pleasant characteristic aroma.
Dried figs can be stored for 6–8 months (Oliveira et al., 2009). There are many
physicochemical quality characteristics of dried fig produced, such as color (O.D. at
340 nm), T.S.S. (%), titratable acidity (T.A. % as citric acid), pH value, rehydration
ratio, and shrinkage (%), which are played an essential role in assessing their quality
and palatability as well as the consumer acceptability of this product.
Various studies have estimated the phytochemical composition in whole fruit of
fresh and dried fig varieties and their skin and pulp fractions. Fresh figs were sub-
jected to two different drying processes: sun drying and oven drying. To assess their
effect on the nutritional and health-related properties of figs, sugars, organic acids,
single phenolics, total phenolics, and antioxidant activity were determined before
and after processing Slatna et al., 2011.
The nutritional profile of dried fig fruit has shown its potential health benefits
(Table 29.1). It has been revealed that the dried fruit of fig has carbohydrates as a
major component (73.5%) that corresponds to its high energy value (317.7 kcal).
Dried fig fruit has a very low amount of fat (0.56%), so it can be helpful for weight
loss. A moderate amount of protein (4.67%) was found in the dried fruit, while
dietary fiber content (3.68%) was good. Figs contain both soluble and insoluble
dietary fiber with many health benefits. Dried figs contained moisture (16.63%) and
high ash content (4.65%). Moisture content affects the texture, taste, appearance,
and stability of foods related to the storage attributes of the dried fruit. The ash con-
tent measures the total amount of minerals present within a food. Mineral content
was also analyzed using ICP-OES.
Dried fig was an excellent source of minerals like Sr, Ca, Mg, P, and Fe (Table
29.2). A relatively high amount of Strontium was found in fig. Strontium has been
found to contribute to good bone health. A patented form of strontium called stron-
tium ranelate is used to treat postmenopausal osteoporosis that reduces the risk of
vertebral and hip fractures. It is the first antiosteoporotic agent that appears to simul-
taneously increase bone formation and decrease bone resorption, thus resulting in
the creation of new bone (Reginster et al., 2007). Calcium is crucial for bones,
29 Fig (Ficus carica) Drying Technologies 679

Table 29.1 The nutritional profile of dried fig fruit (Soni et al., 2014)
Energy Total Dietary
Sample (Kcal/100 g) Carbohydrate Fat Protein fiber Moisture Ash
Dried fig 317.7 73.5 0.56 4.67 3.68 16.63 4.65
fruit

Table 29.2 Mineral content in dried fig fruit (Soni et al., 2014)
Analyte Concentration (ppm)
Sr Saturated
Ca 1545.46
Mg 679.04
P 365.75
Fe 29.49
Zn 9.87
Cu 5.02
Mn 4.75
Sb 0.298
As 1.669
Be N.D.
Cd 0.0034
Cr 1.47
Co 0.032
Pb 0.680
Li 0.245
Mo 0.026
Ni 1.178
Se 0.790
Tl 1.5686
Ti 0.3727
Sn 1.329
N.D not determined

maintaining overall health, and essential for strong bones and teeth. Magnesium is
needed for enzyme action, strong bones and teeth, balanced hormones, and a healthy
nervous and cardiovascular system. Phosphorous is responsible for the growth and
repair of body cells and tissues. Iron is needed for the production of red blood cells
and enzymes (Soni et al., 2014).
The major types of phytochemical compounds in fresh and dried figs include
phenolic acids, flavonoids, and carotenoids. Phenolic acids and flavonoids are the
major types of phytochemical compounds that have been found in both fresh and
dried figs. Their levels are strongly influenced by factors such as the color, the part
of the fruit, maturity, and the drying process. Fresh fig was known to be contained
an adequate amount of antioxidants, and thus, it is necessary to research known a
drying method used led to the excessive retention of antioxidant compounds that
attain the high nutritive and healthy effect of fig products. All tested antioxidants
680 O. Rebai et al.

(L-ascorbic acid, total phenolic, and flavonoid compounds) were significantly

reduced by all drying methods used to determine their original fresh-fig fruit level.
The results indicate that adequately dried figs can be used as a good source of phe-
nolic compounds. Gallic acid, chlorogenic acid, rutin, quercetin-3-O-rutinoside,
and epicatechin are the most predominant phenolic acids and flavonoids in dried
and fresh fig varieties. On the other hand, the literature has reported contradictory
results regarding air- and sun-drying effects on the total content of phytochemical
compounds and the concentrations of individual phenolic compounds and carot-
enoids in figs (Arvaniti et al., 2019). Both fresh and dried figs contain various carot-
enoid compounds; the structures of the most commonly found compounds are
lutein, zeaxanthin, β-cryptoxanthin, α-carotene, β-carotene, and lycopene.

4.3 Nutritional Traits Modification

Many studies have demonstrated that daily intake of fruit and vegetables reduces
chronic-degenerative diseases. In other investigations, it has been observed that
fruit- and vegetable-rich diets protect against different diseases, including cancer
and cardiovascular diseases. Etiology for these diseases points to the free radicals as
promoters of protein, nucleic acids, and cellular lipids oxidations that damage bio-
logical systems; fruits and vegetables contain a significant number of components
with antioxidant activity, such as flavonoids, carotenoids, and vitamins C and E
(Lam et al., 2005).
Furthermore, it has been demonstrated that figs are an essential source of miner-
als and vitamins such as iron, calcium, potassium, thiamin, and riboflavin.
Furthermore, figs are sodium, fat, and cholesterol free; they contain at least 17
amino acids, with aspartic and glutamic acids at the highest concentration. In addi-
tion, fig contains a relatively high fiber content (5.8% w/w), and more than 28% of
it is soluble, so it helps control blood sugar and cholesterol and reduce weight
(Sadhu, 1990).
Fig varieties with dark skin contain high levels of polyphenols, anthocyanin, and
flavonoids, together with higher antioxidant activity than fig varieties of lighter skin
(Solomon et al., 2006). Various plant parts like fruit, root, and figs leaves have
numerous therapeutic benefits. They are used in traditional medicines to treat gas-
trointestinal disorders (colic, indigestion, loss of appetite, and diarrhea), respiratory
disorders (sore throats, coughs, and bronchial problems), and anti-inflammatory
cardiovascular disorders as an anti-inflammatory, antispasmodic remedy. Ficus car-
ica has also been found to have antidiabetic, hypolipidemic, hepatoprotective, anti-
spasmodic antipyretic, antibacterial, antifungal, scavenging activity, and immune
response (Duke et al., 2002).
Today, the fig is a remarkable world crop due to its consumption as fresh or
dry fruit.
The nutritional profiling of the dried fig fruit indicates that it is a good source of
carbohydrates and minerals like strontium, calcium, magnesium, phosphorus, and
iron. It has average protein and dietary fiber content with very low fat. Phytochemistry
29 Fig (Ficus carica) Drying Technologies 681

of the fruit revealed the presence of total phenolics, flavonoids, alkaloids, saponins,
and other secondary metabolites that contribute to its high antioxidant activity (Soni
et al., 2014). Dried figs are high in fiber and carbohydrates and low in fat, making
them healthy food choices. Vitamin A is retained during drying; however, because
vitamin A is light sensitive, food containing it should be stored in dark places. Figs
are one of the highest sources of calcium and fiber in a plant. Furthermore, dried figs
are the richest in fiber and contain essential minerals and vitamins.
The antioxidant activity reported that the antioxidant capacity is frequently due
to phenolic acids, namely gallic acid, caffeic acid, chlorogenic acid, and flavonoids
such as quercetin, catechin rutin, and kaempferol. So far, there are controversial
results in the literature on the role of the drying process on the antioxidant capacity
of figs. In some studies, it has been reported that the high antioxidant capacity is due
to dried figs were higher than those of fresh figs (Slatna et al., 2011; Chang et al.,
2016), while in others, it has been reported that the sun-drying procedure reduced
the antioxidant capacity of figs (Bachir Bey et al., 2016; Kamiloglu et al., 2015).

5 Shelf-Life Extension Using Modified

Atmosphere Packaging

The development of satisfactory methods for the shelf-life extension that ensure
quality maintenance of products with minimum loss has drawn the attention of food
technologists. Modified Atmosphere Packaging MPA or Controlled Atmosphere
Packaging is making a big impact in produce packaging. This advanced process uses
micro-perforation technology to alter the package atmosphere. The ability to effec-
tively manage respiration gasses helps to improve product shelf life considerably.
Over the last years, modified atmosphere packaging (MAP) has received increasing
attention as a method of food preservation. MAP is defined as the enclosure of food
products in gas-barrier materials in which the gaseous environment has been changed
(Sivertsvik et al., 2002). Packaging technologies are important to protect products
against deteriorative effects, including microbial, biochemical, and physical activi-
ties from environmental influences. This involves retardation of spoilage, the exten-
sion of shelf-life, and quality maintenance in packed food. Other packaging functions
include containment, convenience, marketing, and communication (Restuccia et al.,
2010). The rate of deterioration during the storage of a product depends on the bio-
chemical compositions of substrates and metabolites in the tissue, the microbial
contamination, and the condition of storage (Masniyom, 2011).
Modified Atmosphere Packaging is an advanced packaging technology that cre-
ates an optimal environment to extend produce shelf life. Herein, the question is
How Modified Atmosphere Packaging Extends the Shelf Life of the product? The
modified atmosphere created by MAP packaging limits oxygen flow and moisture
evaporation. The controlled gas exchange reduces respiration rates by limiting the
oxygen available in the package. Too much oxygen also increases bacterial decay
and rapid spoilage. Excess oxygen also encourages sprouting, discoloration, and
blemishing. A low oxygen environment produces lower respiration rates. Lower
682 O. Rebai et al.

respiration rates extend shelf life by preventing aging and ensuring moisture reten-
tion in picked produce.
Generally, MAP is used for extending the shelf-life period of fresh or minimally
processed foods. Apart from other perishable products, attempts have been carried
out to prolong the shelf-life of figs by using MAP. It is already known that the fig is
a nutritious fruit, richer in fiber, potassium, calcium, and iron, and is free of sodium,
fat, and cholesterol. Figs also are an essential source of vitamins, amino acids, and
antioxidants. However, the fig is very sensitive to microbial growth even under cold
storage; thus, it is essential to consider alternative processes to extend its shelf life.
Most commercial production is in dried or otherwise processed forms since the ripe
fruit does not transport well and, once picked, does not keep well. The ripening
stages at harvest and post-harvest preservation conditions greatly influence the fig
fruit’s quality (Ayhan & Kara cay, 2011). Modified atmosphere packaging could
provide an alternative method of fruit preservation, longer shelf life, and added
value. Ayhan and Kara cay (2011) showed that considering the sensory, physical,
and chemical qualities, the commercial shelf life of fresh figs is 15 days in the air
and low oxygen applications at 4 °C.
Alturki (2013) examined and monitored the quality parameters to determine the
shelf life of fresh figs under refrigerated Modified Atmosphere Packaging (MAP)
conditions. The fruit quality attributes such as decay, fruit size, external color, firm-
ness, weight loss, water activity, figs fruits' water content, and total soluble solids
were measured during storage at 4 °C for more than 42 days. Modified atmosphere
packaging with 20% CO2 successfully reduced the rate of ripening process while
the control treatment figs showed signs of ripening development such as significant
weight loss and change in figs color. However, the decay incidence was reduced and
delayed by all the MAP treatments. At the end of storage, the weight loss was less
than 1% for all the MAP treatments. The fruit firmness decreased with an increase
in weight loss in each treatment after 28 days of storage. There was no significant
difference in total soluble solids among the MAP treatments. The shelf life of fresh
figs packed in the unsealed cardboard box was less than 14 days.
After storage, all MAP treatments had lost less than 1% of their weight. After
28 days of storage, the fruit firmness reduced as weight loss increased in each condi-
tion. Total soluble solids did not differ significantly across the MAP treatments.
Fresh figs placed in an unsealed cardboard box had less than 14 days of shelf life.
Finally, because MAP improved the external look of the fresh figs, it should be used
commercially in the fresh fig sector for fruit preservation (Alturki, 2013).
Fruits of Ficus carica L. have an exceptional flavor, color, and scent. It does,
however, have a short shelf life due to its morphological peculiarities. Martinez-
Damian et al. (2020) attempt to determine the effect of environment packing on the
nutritional quality and look of figs held at 1 °C. Fruits with decreased weight loss
and firmness were seen using packaging and cold, although citric acid and Vitamin
C levels remained the same. There was no significant fluctuation in the content of
Total Anthocyanins, total phenols, or antioxidant capacity except during the last
evaluation period. On the other hand, a considerable impact on the reduction of
respiration and ethylene generation was seen, which was substantiated by the pres-
ervation of sensory features of the fruit (texture, color, appearance, marketing level,
29 Fig (Ficus carica) Drying Technologies 683

scent, and flavor) from excellent too good. The use of cold (1 °C) and the creation
of passive atmospheres (packing) are effective aids in keeping the nutraceutical
quality and look of fig fruits, both of which are highly valued by consumers and
marketers of this fruit (Martínez-Damián et al., 2020).
In another case, 1-methylcyclopropene (1-MCP) has been frequently used to
improve the shelf life and quality of fruits and vegetables. Researchers discovered
that using paper containing 1-MCP at room temperature can significantly suppress
cell wall degrading enzymes, reduce cell wall polysaccharide disintegration, and
delay softening in Younai plums.
Furthermore, to increase the storage duration of fruits, 1-MCP is frequently com-
bined with other preservation methods such as UV-C irradiation (Pristijono et al.,
2018), calcium chloride (Li et al., 2014), and methyl jasmonate (Ku et al., 2013).
Thus, compared to either therapy alone, a combination of 1-Methylcyclopropene
(1-MCP) and modified atmosphere (MA) treatment has been proven to be more
effective in preserving particular fruits.
The impact of MA and MA+1-MCP treatments on fig fruit postharvest quality
after 30 days at 10.5 °C. Compared to the control and MA groups, the MA+1-MCP
treatment significantly enhanced fruit texture, reduced weight loss and MDA accu-
mulation, and suppressed ethylene production and respiration rate (Fig. 29.3). In
conclusion, the MA+1-MCP treatment will be an effective way to preserve the fruit
quality of figs during postharvest storage.
The color of the peel and flesh is the most evident and dependable indicator for
customers to determine the quality of fruits and vegetables. Figure 29.3 shows that
after 30 days of storage, the fig fruit had varying degrees of degradation. The flesh

Fig. 29.3 Effects of MA and MA + 1-MCP treatments on deterioration development in fig fruit
after storage for 30 days at −1 °C: (a) fresh fruits; (b) MA treatment and stored for 30 days; (c)
MA + 1-MCP treatment and stored for 30 days; (d) the control fruit stored for 30 days (Song et
al., 2019)
684 O. Rebai et al.

of MA+1-MCP-treated fruits is superior to that of MA-treated and control fruits.

The 2e flesh of MA+1-MCP-treated fruits is the best, whereas control fruits are
nearly inedible.
In conclusion, MAP resulted in a better external appearance, and therefore the
commercial application of MAP in the fresh fig industry should be introduced for
fruit preservation.

6 Conclusions and Future Prospects of Fig Drying

There is a large amount of research, and available knowledge, carried out by aca-
demia, but the industry is not taking full advantage even in the more developed coun-
tries. A lack of interaction between researchers and industry is observed worldwide;
consequently, industrial advancement is slow because of a mismatch between
research and industrial needs. In less developed countries where the industry is not
very important, there is a general feeling that drying is an easy operation and not too
much input is needed, and anybody can do it. A consequence of this attitude has been
the failure of many drying projects. Drying foods in general and figs, in particular, is
a complex business, and a mere translation from other fields is not often advisable.
Energy efficiency linked to environmentally friendly processes and products also
appear as a growing trend, mainly in some developed countries. Therefore, there is
a need to check existing data on drying kinetics to establish their applicability
regarding the prevailing resistance. Indeed, there is a need to study products (iso-
therms and drying kinetics) not common in more developed countries where the
leading research in the field has been carried out. This will lead to discovering new
niches of activity. This is also true for countries where drying research is an active
field where the use of off-quality fresh market products is often disregarded. One
can say that there is a general concern about improving the final consumer products
quality. Consequently, pretreatments (including enrichment), optimization and con-
trol of operating conditions, and a combination of fig drying strategies according to
the drying stage should be studied better to migrate water, storage, and nutritional
profile consumption.
Though good feeds have been achieved in drying technology, there is a need to
develop cost-effective and energy-efficient drying methods further. The desirable
aspects of fruits and vegetables need to be preserved during the drying process.
Combinations of drying techniques produce products that are more appreciated by
the consumers and reduce the heat-induced deterioration in fruits and vegetables. On
the other hand, the consumption of high energy by instruments used for drying adds
to the cost of the finished end product. The application of the natural drying method
reduces the cost and is much easier to use. The major drawback of the natural drying
method is that it is time-consuming, and the drying conditions cannot be manipu-
lated compared to the artificial drying method. Future work could focus on develop-
ing natural drying solutions with greater efficiency. Drying kinetics has helped
achieve good and effective results with minimal resource utilization. However, dry-
ing models that consider all the variables are still non-existing, making it a potential
29 Fig (Ficus carica) Drying Technologies 685

research area. Advances in dehydration techniques and the development of novel

drying methods have enabled the preparation of a wide range of dehydrated products
and convenience foods from fruits and vegetables, meeting the quality, stability, and
functional requirements coupled with the economy. This has been made possible by
sustained experimental studies to understand the theoretical and fundamental aspects
of the process and optimization of the techniques to achieve a favorable combination
of cost and quality. This paper reviews and highlights the developments of the most
popular drying methods of fig and their products during the last decade covering
theoretical aspects and practical applications with major emphasis on techniques
that have received the maximum attention (Ahmed et al., 2013).

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Chapter 30
Chemistry and Functionality of Processed
Figs

Asad Nawaz, Noman Walayat, Ali Hassan, Maryam Chaudhary,

and Ibrahim Khalifa

1 Introduction

Figs are widespread dry fruits usually grown in warm and dry climates. The best
condition for intensive figs cultivation is a semi-arid climate and irrigation. The figs
production worldwide is more than one million tons, and it is most common in the
Mediterranean region. In this region, figs have been cultivated for centuries. The
trees of figs produce one or two crops yearly, depending on the cultivar type. The
primary crop is grownup from flowers introduced in the previous year, and the fruit
gets ripened at the start of summer. The second main crop is produced from flowers
that appear in the present season, and the fruit gets ripened in late summer. Hence,
the growth of both crops is evident in different climatic conditions. Fruits cultivated
from the two crops may also vary in shape and size. Figs are commonly consumed
fresh and also peeled in dried form. Naturally, fresh fruits have a short post-harvest
life span of 7–10 days, but a combination of favourable conditions and carbon diox-
ide in the atmosphere enhances the fruit shelf life and can be stored for up to

A. Nawaz
Shenzhen Key Laboratory of Marine Microbiome Engineering, Institute for Advanced Study,
Shenzhen University, Shenzhen, China
N. Walayat
College of Food Science and Technology, Zhejiang University of Technology,
Hangzhou, China
A. Hassan · M. Chaudhary
National Institute of Food Science and Technology, University of Agriculture,
Faisalabad, Pakistan
I. Khalifa (*)
Food Technology Department, Faculty of Agriculture, Benha University, Moshtohor, Egypt
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 689
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_30
690 A. Nawaz et al.

2–4 weeks. Figs are also commonly consumed as dried fruit, as the drying method
extends their storability and maintains quality characteristics (Veberic et al., 2008).
Figs are used in primary and processed forms to produce traditional and indus-
trial products like jams, infusions, wines, liqueurs, spirits, etc. Fig fruit is a perish-
able food item, primarily consumed in dried form. Fresh or dry figs are considered
very important for their high nutritive value, high mineral content, fats, sugars, cal-
cium, and presence of other non-nutritive elements such as fiber, water, and antioxi-
dants like phenolic compounds that pose numerous health benefits. In addition, the
decoctions, infusions, and processing of fig leaves have traditionally been used to
treat diseases because of their therapeutic effect and chemical characteristics
(FR, 2013).
The processing of figs in dried form is nutritious with high potential benefits. In
addition, dried fig is a rich source of bioactive compounds with health-promoting
benefits. However, dried figs’chemistry and quality characteristics depend on differ-
ent factors like harvesting practices, stages of ripening, geographical region, and
processing techniques. Before harvesting figs, the fruits are left on the tree until they
get fully ripe, dried partially, and fall to the ground. Then, semi-dried figs are placed
on trays made of wood for further drying and kept for 2–3 days to achieve 22–24%
dry matter. Finally, these dried figs are transferred to industries for further process-
ing to increase and maintain the final product quality. The processing steps involved
in drying figs are sorting, fumigation, cleaning, washing, proper shaping, and final
packaging.
It is essential to process or wash figs under controlled conditions as different
treatments during washing of figs may affect the textural, sensorial, chemical, and
color properties and may lead to loss of texture, flavor, and essential nutrients of the
final product. Furthermore, the washing of figs is helpful in the removal of pesti-
cides or microorganisms, therefore minimizing the adverse effects on human health.
The most used technique for washing figs in food industries is with chlorinated
water (using sodium chloride) because it is cost-effective and efficiently decontami-
nates the microorganisms. Although chlorine is used as a popular sanitizing agent,
improper usage may cause deterioration of quality by reacting with other organic
compounds present in food items. Therefore, it is imperative to watch or monitor the
free level of chlorine, as if it is not monitored properly, it may lead to the generation
of carcinogenic by-products. Recently, different techniques such as ultrasound and
pulse electric field have been used as an alternative treatment to chemicals used dur-
ing washing. The most promising technology is the US technique which is non-
thermal technology as it led towards microbial safety and shelf-life extension. It
also retains the nutritional value, sensory, chemical, and functionality of food items.
Additionally, this technique has less use of the chemical sanitizing agent, mini-
mal human contact, and consistent and uniform cleaning. However, the US tech-
nique alone is not sufficient for proper surface decontamination. Therefore, different
studies suggested the combined use of the US technique and sanitizing agents in
washing during the processing of dried figs to retain their chemical and physical
properties (Gençdağ et al., 2021).
30 Chemistry and Functionality of Processed Figs 691

2 Types of Processed Figs

One of the Mediterranean basin’s oldest fruit species, the common fig may produce
two harvests every year. However, the quality of the crops may vary regarding sea-
son, variety, and harvesting stage (Pereira et al., 2017). Figs may be consumed
fresh, dried, or processed into various goods, including jams, jelly, squashes,
smoothies, wines, and bakeries (Barolo et al., 2014). Generally, processed figs are
consumed compared to fresh ones due to the high moisture content in fresh fruits
(Teruel et al., 2021). Fruit drying is the simplest method of preserving fig fruit that
is usually processed utilizing some unit operations, such as drying and freezing
(Yao et al., 2021). The conventional sun-drying procedure produces a wide range of
quality food owing to its ability to retain phenolic and flavonoids compound with
minimum loss of nutrition (Manoj et al., 2018).
Meanwhile, automated air dehydration offers various benefits, including produc-
ing fruit with high sugar content and total phenolics. More recently, the freeze-
drying method has widely been used for the drying of figs in order to preserve their
original phenolic compounds and bioactive compounds (Martínez et al., 2013).
Besides this, the processing of figs through fermentation has also been reported to
retain a high amount of phenolic and bioactive compounds (Buenrostro-Figueroa
et al., 2017). Interestingly, fig fruits have been frozen at −18 °C to preserve their
nutritional content and make the best use of figs in off seasons (Petkova et al., 2019).
Another study (Palmeira et al., 2019) also reported the freeze-drying of fig fruit
parts (peel and pulp) at −80 °C and found that antioxidant and phenolic compounds
were higher in the peel as compared to the pulp.
Similarly, another study (Bachir bey et al., 2014) also reported the freeze-drying
of a dark-purple skin and red pulp to preserve the phenolic compounds. Thus, the
freeze-drying method is widely used to preserve the phenolic and bioactive com-
pounds. Furthermore, because fresh or dried fig fruits have a high concentration of
beneficial chemicals, their use should be promoted as a possible healthy substitute
for sweets. Thus, the right choice of processing is preferred to maintain its native
nutrients.
Besides processing, fig fruits have been utilized in numerous processing foods,
such as biscuits, jam, jelly, confectionery, soups, etc. (Teruel et al., 2021). It has
been established that the use of fig fruits in different products resulted in a higher
nutritional value and the accumulation of various bioactive compounds. For
instance, the study reported that the addition of fig seed powder in bakery products,
such as buns and muffins, increased fiber content and total phenolic and antioxidant
activity (Chauhan, 2016). Another study reported the addition of fig seed powder
replacing wheat flour in biscuits, which resulted in increased protein and fiber con-
tent and an upsurge in total phenolic compounds (Bölek, 2021). In addition, the use
of fruits in desserts and smoothies has been encouraged or reported as an alternative
to sugar, along with the increased phytochemicals (Jahromi & Niakousari, 2018).
However, in some studies, the processed figs, especially the dried ones, resulted in
lower total phenolic compounds in wines when dried figs were added compared to
692 A. Nawaz et al.

the fresh figs (Lu et al., 2021). In conclusion, the use of fig fruits to develop func-
tional food has potential and practical significance.

3 Composition of Processed Figs

Figs have remained the main part of the diet and symbol of health in Middle East
Countries. However, a complete understanding of the composition of processed figs
is inevitable. Fresh and dried figs are high in fiber, amino acids (aspartic acid and
glutamine), vitamins (thiamine, riboflavin), carotenoids (lutein, cryptoxanthin,
lycopene, -carotene), minerals (iron, calcium, potassium), antioxidant polyphenols,
sugars, and organic acids (Chauhan, 2016). Furthermore, figs are low in sodium, fat,
and cholesterol. These Ficus species have been shown to have positive health-
related effects due to their phytochemical composition, primarily flavonoids, which
appears promising in lowering blood lipid levels and preventing conditions such as
obesity, diabetes, cardiovascular disease, neurodegenerative disorders, and even
certain types of cancer (Caliskan, 2015; Wojdyło et al., 2016). It is understood that
the processing of figs results in compositional changes that may be beneficial, or
sometimes they losses nutritional values. Usually, figs are processed to increase
their shelf life and nutritional values. Because of their high sugar concentration and
low organic acid level, they have a sweet flavor. Their phenolic content is moderate,
with red varieties containing larger quantities of anthocyanins. Compared to other
fruits, the fig has low total carotenoids, concentrated mainly in the fruit’s skin
(Pereira et al., 2017). Table 30.1 shows the antioxidant capacity, total phenols, and
total flavonoids of various types of processed figs. Several studies have reported that
fig fruit has multiple health benefits, including antibacterial, antidiabetic, anti-
inflammatory, and anticancer (Caliskan, 2015; Li et al., 2021). However, the pro-
cessing of figs, because of heat treatment, may cause a reduction in these
characteristics, such as loss of phenolic and flavonoid compounds.
Arvaniti et al. (2019) reported the composition of fresh and dried figs, and results
showed that protein fat and total sugars were found to be highest in dried figs
(3.30%, 0.93%, and 47.9%, respectively) as compared to fresh one (0.75, 0.30 and
16.2, respectively), whereas water was highest (79.3%) in fresh one compared to the
dry one. Moreover, the study reported that dried fig fruits had ample amounts of
minerals (Na, K, P, Zn, Mg, Fe, and Ca) and vitamins (A, C, B1, and B2). Petkova
et al. (2019) studied the composition of figs fruits in fresh and frozen storage
(−18 °C for 4 months) and reported that moisture and pH were increased in frozen
storage, whereas ash content was decreased during frozen storage. Slatnar et al.
(2011) studied the composition of processed figs dried by sun drying and oven dry-
ing. The results showed fructose (~52%) was highest among the sugars, followed by
glucose (~46%) and sucrose (~2%). Interestingly, it was found that the oven drying
method resulted in a high number of total sugars compared to sun-drying and fresh.
Another study found higher total sugar content in fresh figs. (23.5 g/100 fw) as
compared to frozen figs. (19.2 g/100 g fw) (Petkova et al., 2019). Similarly,
30 Chemistry and Functionality of Processed Figs 693

Table 30.1 Total phenols, flavonoids, and antioxidant capacities of processed figs
Types/
variety of Processing Total Antioxidant
figs method Total phenolics flavonoids capacity References
Bela Drying oven 530 mg.GAE N.D N.D Slatnar et al.
petrovka kg−1 dw (2011)
Dry figs N.A 195.33 mg/100 g N.D 0.388 mmol/100 g Miletić et al.
from Serbia dw dw (2014)
Dried figs Oven drying 19.2 mg/100 g N.D N.D Vallejo et al.
from fw (2012)
Turkey
Dried Sun-drying 201.7 mg 112.28 mg 131.55 mg Khadhraoui
Saoudi (GAE/100 dw) (QE/100 g (TEAC/100 g dw) et al. (2019)
Douiret dw)
Mission Freeze drying 3.08 mg CE/g N.D 2.0 μM eq Martínez
dried figs Trolox/g et al. (2013)
Mission Drying at 3.23 mg CE/g N.D 3.7 μM eq Martínez
dried figs 55 °C Trolox/g et al. (2013)
Mission Drying at 3.72 mg CE/g N.D 3.83.7 μM eq Arvaniti
dried figs 65 °C Trolox/g et al. (2019)
Fermented Fungal 4.77 mg GAE/g. N.D 0.53 mg of GAE/g Buenrostro-
figs strains dw of dm Figueroa
Rhizopus et al. (2017)
oryzae at
30 °C for
5 days
Cuello 60 °C during 2.67 mg GAE/g 17.61 mg DPPH 9.50%/g Viuda-
dama 24 h RE/g Martos et al.
Powder figs (2015)
(peel)
N.D means not detected, dw means dry weight

regarding organic acids, it was found that malic acid (25–59% of total organic acids)
was highest compared to citric acid, while it was highest in processed figs compared
to fresh fruits. Pande and Akoh (2010) also reported the highest malic acid content
in figs, especially in peel and pulp, compared to whole fruits. Regarding phenolic
compounds, it was found that figs contain many phenolic compounds, including
hydroxycinnamic acids, flavan-3-ols, flavanols, and anthocyanins. However, it has
been reported that crop season also differs the phenolic contents. In addition to this,
it has been found that processing methods significantly affect the composition of
phenolic compounds (Slatnar et al., 2011), lower in fresh and higher in processed
figs, especially oven drying compared to sun drying. It is believed that oven drying
results in the loss of phenolic compounds, but recently, it has been revealed that they
are better preserved by the drying process than hydroxycinnamic acids or mono-
meric catechin (Devic et al., 2010). This is because monomeric catechins are
responsible for enzymatic browning, but they are also easily diffused due to their
lower molecular weight. On the other hand, Arvaniti et al. (2019); Slatnar et al.
694 A. Nawaz et al.

(2011) reported the degradation of phenolic compounds in dried figs. Arvaniti et al.
(2019) also found the loss of 15% of total phenolic compounds due to drying com-
pared to a fresh one in figs “Cuello Dama”.
Besides this, safety is the major concern during the processing of figs. Myotoxicity
and mould growth are the major toxins that may produce during processing. Iqbal
et al. (2018) reported the presence of aflatoxin and ochratoxin in dried figs. Similarly,
another study also reported the presence of aflatoxin in dried figs (Petrić et al.,
2018). However, pre-treatments can help reduce these toxins, an important safety
concern. Several types of food additives, such as citric acid, emulsifiers, amino
acids (e.g., L-cysteine), and antioxidants, may help reduce toxins.

4 Bioactive Components of Processed Fig

Fig fruits have excellent sources of bioactive compounds reported to cure various
diseases, such as atopic dermatitis, skin warts, and cervical cancer. Teruel et al.
(2021) comprehensively studied a detailed meta-analysis of by-products of fig fruits
and found 19, 21, and 22 types of phenolic compounds in whole fruits, peel, and
pulp, respectively. Moreover, these compounds were from various families, such as
flavonoids, phytosterols, coumarins, hydroxycinnamic acids, anthocyanins, and
phenolic acids. Slavin (2006) reported that figs are the richest source of benzalde-
hyde used to cure skin diseases. Weibin et al. (2001) reported that dried figs, espe-
cially by sun-drying, contained bioactive compounds, such as flavone, routine, and
quercetin used for the formulation of medicines used for curing cardiovascular dis-
ease. It has been found that figs are excellent sources of Laxatives bioactive com-
pounds that help increase stool motility and reduce constipation (Tabrizi et al., 2020).
Regarding the processing of individual parts, it was found that peel contained
more phenolic compounds as compared to the pulp of freeze-dried fig fruits. The
study found 18 phenolic compounds in fig peel of green fig, among which 8 were
flavonoids, and the remaining were phenolic acids and their derivatives (Palmeira
et al., 2019). The same study reported that caffeic and vanillic acid derivatives were
abundant among the phenolic acids, whereas quercetin and C-glycosylated apigenin
derivatives were the most abundant flavonoid compounds. Similar results were
reported by another study when peel and pulp were evaluated for phenolic com-
pounds. Agreeing with these findings, Harzallah et al. (2016) also reported the more
phenolic compounds in the peel of three varieties (green, purple and black figs);
however, the black variety showed more antioxidant capacities. Therefore, it is
inferred that bioactive compounds differ greatly in varieties having darker colors
will have more phenolic and flavonoid compounds. These compounds also depend
on the ripening and/or harvesting stages: more ripened fruits have more phenolic
compounds than the unripen ones.
30 Chemistry and Functionality of Processed Figs 695

5 Stability of Bioactive Components

5.1 Bioactive Components

“Bioactive components play an important role in the body and help to promote good
health. This term refers to non-essential biomolecules in foods that exhibit the
potential to modulate metabolic processes and different mechanisms in the human
body to optimize human health”.
Plants secondary metabolites or phytochemicals are non-nutritive metabolites of
plants that are important for plant growth, survival, and reproduction. Most of these
components are biologically active compounds and provide health benefits. F. car-
ica plants exhibit the highest number of compounds. Phytochemical studies showed
the presence of bioactive compounds and other secondary metabolites in figs plant
(Barolo et al., 2014). Fig fruits have been consumed primarily in dried and fresh
form; they have also traditionally been processed and preserved. Nowadays, con-
sumers demand products based on fig fruit because of their high nutritional profile
and different bioactive components in figs (Slavin, 2006).
Figs comprise many bioactive compounds in the flesh, peel, leaves, and whole
fruits. The bioactive components present in figs are chlorogenic acid, cyaniding,
luteolin, rutin, and catechin. These bioactive compounds posed potential health ben-
efits such as hepatoprotective, antibacterial, antidiabetic, antioxidant, anti-
inflammatory, and anticancer properties. Therefore, the demand for figs and
fig-based products has increased by consumers.
The bioactive components present in whole figs are catechin, epicatechin, poly-
meric procyanidins, rutin, chlorogenic acid, cyanidin-3,5-O-diglycoside, cyanidin-3-
O-rutinoside, pelargonidin-3-O-rutinoside, kaempferol-3-rutinoside,
kaempferol-3-glucoside, quercetin-3-glucoside, quercetin-3-O-rutioside,
quercetin-3-galactoside, quercetin-3-O-malonyl-galactoside, apigenin-C-hexoside-
pentoside, gallic acid, syringic acid, and ellagic acid. In whole figs, Quercetin-3-O-
rutioside is considered the major individual phenolic compound followed by
quercentin-3-glucoside, chlorogenic acid, and polymeric procyanidins cyaniding-3-
O-rutinoside. The most abundant bioactive compounds considered in the peel are
cyanidin-3-O-rutinoside followed by cyanidin-3-O-diglucoside, cyanidin-3,5-
diglucoside, catechin, epicatechin, and quercetin-rutinoside. Cyanidin-3-rutinoside
and epicatechin are the main bioactive compounds in pulp of fig, whereas caftaric
acid in the form of kaempferol-3-O-glucoside is considered to be the main compo-
nent in leaves of figs (Teruel et al., 2021).
Contents of the bioactive compound and antioxidant activity in figs depend on
the type of cultivator in both processed and fresh figs. Fresh and dried figs are a
good source of carbohydrates, amino acids, fiber, sugars, vitamins, minerals (man-
ganese, copper, magnesium, calcium, and potassium), phenolic compounds, and
696 A. Nawaz et al.

organic acids. Phytochemicals present in figs are arabinose, β-carotenes, β-amyrins,

β-sitosterols, glycosides, xanthotoxol, flavonoids, alkaloids, saponins, coumarins,
and terpenes.
There is an impact of different processing conditions on the bioactive properties
of figs. The research was conducted by some researchers who evaluated phytochem-
ical components in frozen, fresh, and processed figs. Proximate analysis and total
chlorophyll, total phenols, total carotenoids, total anthocyanins, and carbohydrates
contents were evaluated. In addition, they determined the antioxidant content of
frozen, fresh, and processed figs by FRAP and DPPH· assays. The study’s findings
depicted a high range of antioxidant activity and bioactive components in fresh fruit
compared to frozen fruits in which the levels of bioactive components decreased
significantly. To preserve bioactive components, mainly phenolic and carotenoids,
processing figs in the form of jams is the best option. However, after a few months
of storage at a temperature of −18 °C, significant losses in bioactive compounds
were observed. Regardless, figs processing in the form of the jam was considered to
retain a better amount of bioactive compounds than storage in freezing (Petkova
et al., 2019).
The antioxidant activity and bioactive compound content depend on the type of
cultivator in both processed and fresh figs. One of the research projects was con-
ducted in which main phenolic compounds were studied in nine cultivars of sun-
dried figs with different skin colors. For all parameters, a considerable difference
was observed between the cultivars. Dark-colored cultivars contained a high level of
phenolic and flavonoids and high oxidant potential than light-colored cultivars
(Khadhraoui et al., 2019).
One of the studies was conducted in which the impact of figs processing on nutri-
tional, physicochemical, and phytochemical properties was investigated. They
washed the figs and performed the other operations to process them into jam or
nectar. The physicochemical parameters, iron, phosphorus, and calcium content
were evaluated. Proximate analysis, vitamin content, beta-carotene, and total carbo-
hydrates were also analyzed for nutritional analysis after processing. Different phy-
tochemicals determined were total flavonoids, total phenolic, anthocyanins, and
tannins. The findings of this study depicted that the processing of fig pulp into nec-
tar and jam caused a significant increase in physicochemical properties, but a sig-
nificant decrease in iron, pH, phosphorus, and calcium was observed. Processing
the pulp of fig into jam and nectar caused a decrease in nutritional properties and
phytochemical composition compared to fig pulp. Proximate composition, vitamin
C, β-carotene, total flavonoids, total phenolic, tannins, and total anthocyanins
decreased significantly due to processing, whereas a significant increase in carbohy-
drate was observed that caused an increase in energy value (Tanwar et al., 2014).
30 Chemistry and Functionality of Processed Figs 697

6 Functionality of Processed Fig

6.1 Functional Properties

Food’s functional characteristics are the important physicochemical properties that

reflect the interactions between the molecular compositions, conformation, struc-
tures, and physicochemical properties of different components present in the food
according to the environmental conditions in which they are associated and mea-
sured. Functional properties are required to predict and evaluate how fats, proteins,
carbohydrates, and fiber act on specific food items. Functional characteristics also
define the action of ingredients during processing, preparation, and cooking and
their effect on the final product after processing, depending on its taste, appearance,
and texture. Food functionality is characterized by food products’ quality, structure,
nutritional composition, appearance, and acceptability (Awuchi et al., 2019)
(Table 30.2).
The proper processing and storage of figs are required to retain their nutritional
and functional properties, as figs are very sensitive to microbial spoilage or growth
under cold storage conditions. Therefore, it is essential to use another alternative
process to increase their shelf life. The processing of figs, including the drying
method, causes many changes, such as the size of figs, compositional properties,
and thermodynamics parameters of drying air such as humidity, temperature, and
velocity. The volume and mass of figs get reduced by the drying method, which also

Table 30.2 Nutritional composition of figs

Dietary element Fresh fig (per 100 g) Dried fig (per 100 g)
Total calories (kcal) 74.0 249.0
Water (grams) 79.11 30.05
Protein (grams) 0.75 3.30
Total fat (grams) 0.30 0.93
Saturated fat (grams) 0.06 0.93
Sugars (grams) 16.26 47.92
Fiber (grams) 2.9 9.8
Calcium (milligrams) 35.0 162
Cholesterol (milligrams) 0 0
Phosphorus (milligrams) 14.0 67.0
Magnesium (milligrams) 17.0 68.0
Iron (milligrams) 0.37 2.03
Potassium (milligrams) 232 680
Zinc (milligrams) 0.15 0.55
Vitamin C (milligrams) 2.0 1.2
Vitamin A (IU) 142.0 10
Riboflavin (milligrams) 0.050 0.085
Thiamin (milligrams) 0.060 0.085
Source: USDA National Nutrient Data Base for Standard References Haytowitz et al. (2018)
698 A. Nawaz et al.

affects the handling and packaging cost. The drying process also causes a reduction
in microbial activity and retains the chemical properties of the final product
(Martínez et al., 2013).

6.2 Fig as a Functional Food

Functional foods are “foods containing physiologically active compounds that pro-
vide benefits along with basic nutrition and prevent us from diseases or promote
health”.
Figs are harvested worldwide and consumed in fresh and dry forms. Its edible
portion is the fruit, hollow, fleshy, and receptacle. Figs are an essential source of
minerals, vitamins, carbohydrates, organic acids, sugars, and phenolic compounds.
Dried, fresh, and processed figs contain high amounts of polyphenols and fiber. Figs
have a high concentration of phenolic compounds such as proanthocyanidins. The
phenolic compounds present in figs are higher in content than tea and red wine,
which are good sources of phenolic compounds. Fig roots, fruit, and leaves are used
in traditional medicines to treat different diseases such as gastrointestinal, cardio-
vascular, and respiratory disorders and are used as antispasmodic and anti-
inflammatory remedies (Mawa et al., 2013).
Fig is considered a remarkable crop globally because of its consumption on a dry
and fresh basis. Fig fruit is an essential source of vitamins and minerals such as
calcium, potassium, iron, riboflavin, and thiamine. Figs are rich in fat, sodium, and
cholesterol-free and provide the body with all the vital nutrients needed. Figs con-
tain nearly 17 essential amino acids, of which glutamic and aspartic acid are present
in the highest concentration. The high fiber content is present in figs, and most of the
fiber present is soluble. Therefore, it helps regulate blood sugar levels, improve
cholesterol level, and help maintain a healthy weight. The concentration of polyphe-
nols is high in dry figs compared to other beverages and fruits. Because of these
important health benefits, figs are considered an important functional food (Solomon
et al., 2006).
Antioxidant compounds such as vitamin E, phenolic, organic acids, and carot-
enoids scavenge free radicals and inhibit the oxidative mechanism that leads to
degenerative disorders. Phenolic compounds are common secondary metabolites of
plants that provide physiological and health benefits to human health because of
their antioxidant properties. Phenolic compounds help to regulate body mechanisms
by reducing or donating hydrogen atoms, acting as free radical scavengers. These
phenolic compounds are essential components of the flavor, color, and aroma of
fresh vegetables, fruits, and other food products. In addition, phenolic compounds
also play anti-inflammatory, anti-mutagenic, anti-carcinogenic, and exhibit antimi-
crobial properties.
F. carica comprises a high level of antioxidants and phenolic compounds that
play an important role in promoting human health. These phenolic compounds act
as antioxidants in different ways, as hydrogen donators, reducing agents, free
30 Chemistry and Functionality of Processed Figs 699

radicals scavenging, and singlet oxygen quenchers. High levels of flavonoids, poly-
phenols, antioxidant potential, and anthocyanins’profile in figs help treat many dis-
eases. The antioxidant potential in figs may help protect lipoproteins in plasma from
oxidation and enhance the antioxidant capacity of plasma. Figs are high in essential
minerals required to optimize body mechanisms. Figs also consist of sugars, mainly
glucose and fructose. Anthocyanin content in different parts of figs helps to regulate
body metabolism (Çalişkan & Polat, 2011). A combination of compounds 6-O-acyl-
β-d-glucosyl-β-sitosterols in figs was found to have an inhibitory effect on the pro-
liferation of cancer (Rubnov et al., 2001).
In many countries, figs have been used as a therapeutic medicine. Figs plant
exhibits antioxidant, antifungal, anti-carcinogenic and anti-helminthic activities and
inhibits acetylcholinesterase. Figs are also considered to be effective against
oesophageal cancer cell lines. It also exhibits antifungal and anti-bacterial proper-
ties. Leaf extracts of figs increased the protection against CCl4 hepatic damage and
are considered a hepatoprotective. It also exhibits antipyretic potential. Figs are also
considered anti-hypertensive and reduce coronary heart disease in hypertensive
patients. Bioactive compounds in figs increase insulin secretion and reduce blood
glucose and have been anti-diabetic (Hashemi & Abediankenari, 2013).
Flavonoids, alkaloids, coumarins, terpenes, and saponins are found in an aque-
ous extract of dried ripe fruit of F. carica. Phenolic compounds isolated from the
leaves of figs are reported to exhibit pharmacological properties. These compounds
have diverse biological mechanisms and provide health benefits. The extract of
methanol from F. carica is effective against the activity of oral bacteria and acts as
an anti-bacterial agent. The combination of methanol with gentamicin and ampicil-
lin was found to be synergistic against oral bacteria, which shows that figs act as
natural antibacterial agents (Jeong et al., 2009).

7 The General Impact of Figs on Health

Figs are considered very important for the proper functionality of the human body.
Some important benefits of figs are summarized below.
• Figs satisfy the cravings for sweets and are considered a healthy alternative com-
pared to other sweet food items and can be taken as a snack and used in dairy
products.
• Figs are rich in potassium and help overcome potassium imbalance in the body.
High sodium levels in figs help maintain sodium balance in the body. Therefore,
figs also help maintain blood pressure by maintaining a sodium potassium imbal-
ance in the body.
• Figs also help improve digestive issues, prevent diarrhea, constipation, and aid in
digestion.
• Figs are considered a good source of prebiotics that improves overall gut health.
700 A. Nawaz et al.

• Figs help increase bone density as they are an excellent source of potassium and
calcium. These essential minerals help to improve bone health and prevent
osteoporosis.
• Figs are rich in bioactive compounds and polyphenols that act as antioxidants in
the body and prevent cancer-producing cells.

8 Conclusion

In this chapter, the processing, chemistry, and functionality of fig fruits have been
discussed. Various processing methods have been discussed, by which freeze-drying
method is the best choice to retain its bioactive compounds. On the other hand, the
oven drying method has been proposed as an alternative to sun-drying due to safety
concerns, e.g., aflatoxins. Regarding composition, it has been established that peel
contains more phenolic and bioactive compounds than seeds and/or whole fruits. In
addition, varieties, such as dark-colored fruits, have more phenolic and bioactive
compounds than green whereas ripen fruits have more sugars and flavonoids than
unripe and green ones. The bioactive compounds in processed figs have the poten-
tial to cure numerous disorders, such as skin problems, digestion disorders, cancers,
and heart diseases. Furthermore, processed figs have the potential to be used in
numerous food products, which can be served as functional foods.

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Chapter 31
Fig (Ficus carica) Syrup as a Natural
Sugar Substitute

Akram Sharifi , Elham Taghavi , and Sara Khoshnoudi-Nia

1 Fig and Fig Compounds

Fig (Ficus carica L.) is an important crop worldwide and one of the most abundant
fruits in the Mediterranean diet for dry and fresh consumption. Fig industrialization
produces many co-products. These co-products come from fresh figs discarded
because of inadequate ripening or over-ripening, spoiled, size, texture, or low qual-
ity as table fruit, but they are safe for human consumption. The non-use of fig co-
product constitutes an economic loss since it is rich in nutrients and bioactive
compounds, which can be extracted and used as value-added materials. In addition,
figs and figs co-products are an excellent source of minerals, vitamins, and dietary
fiber; they are fat and cholesterol-free and contain many amino acids (Viuda-Martos
et al., 2015).
Fresh fig is well known for its attractive taste and excellent nutritional composi-
tion. In addition, it is a highly nutritious fruit and is rich in calories and carbohy-
drates, lipids, phenolics, and enzymes. Figs are known to contain numerous phenolic
compounds with potential antioxidative activity (Veberic et al., 2008). Sugars,

A. Sharifi (*)
Department of Food Science and Technology, Faculty of Industrial and Mechanical
Engineering, Qazvin Branch, Islamic Azad University, Qazvin, Iran
e-mail: [email protected]
E. Taghavi
Faculty of Fisheries and Food Science, Universiti Malaysia Terengganu,
Kuala Nerus, Terengganu, Malaysia
e-mail: [email protected]
S. Khoshnoudi-Nia
Seafood Processing Research Group, School of Agriculture, Shiraz University, Shiraz, Iran
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 703
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_31
704 A. Sharifi et al.

acids, and phenolic compoundscontribute to the taste and color of figs and the flavor
characteristic, which mainly depends on the proper balance of the volatile chemical
constituents (Pereira et al., 2020). For example, 100 g of fresh Fig (Ficus carica L.)
contains 86% water, 310 kJ energy, 19.2 g carbohydrate, 16.3 g sugars, 0.3 g total
fat, 0.7 g protein, and T.S.S.(°Bx) = 18. (Hussain et al., 2021; Veberic & Mikulic-
Petkovsek, 2016; Pawase et al., 2017). Vitamins and minerals of fresh fig (Ficus
carica L.) fruit are shown in Table 31.1.

1.1 Carbohydrate

Fig is a high carbohydrate fruit and a good source of dietary fiber. The carbohydrate
content of fresh fig is 19.2 g/100 g of fresh fruit weight. About 92% of the carbohy-
drates in figs are in the form of sugars. Rest consists of dietary fiber, insoluble cel-
lulose in the skin, and soluble protein in fruit. The total sugar content of fig is
16.3 g/100 g, while dietary fiber constitutes about 5 g/100 g. Some fig varieties
contain β-D glucans with substantial antimicrobial activity (Hussain et al., 2021).
Glucose, fructose, and sucrose are the main sugars present in fig fruits. Sorbitol
is present at low concentrations, and, therefore, the fig is considered a sorbitol-poor
species (Mascellani et al., 2021). The content levels of glucose, fructose, and sucrose
are 1216–11,050 mg/100 g, 1916–11,900 mg/100 g, and 98–320 mg/100 g, respec-
tively. Only traces of sucrose have been measured in analyzed fig fruit and juice
extracted from figs. Çalişkan and Polat (2011) showed sucrose content in 76 local
Mediterranean fig accessions. In some accessions, sucrose was not determined; the
highest value was measured in ‘Mor 3’ (320 mg/kg). Other authors also report that
this sugar is not present in fig juice or not distinguishable from other sugars (Çalişkan
& Polat, 2011). Viuda-Martos et al. (2015) showed sucrose, fructose, and glucose in
fig powder 5.91, 67.3, and 68.2 (mg/g sample).
In contrast, sucrose is the prevalent sugar in fig leaves. Sucrose is the main trans-
location carbohydrate from leaves to other plant parts, including fruit. The low
sucrose concentration in fig fruit could result from anabolic processes and

Table 31.1 Vitamins and minerals of fresh fig (Ficus carica L.) (Hussain et al., 2021)
Component Nutritional value/100 g Component Nutritional value/100 g
Vitamin A Equiv. 142 IU Vitamin E 0.1 mg
Thiamine (B1) 0.1 mg Phosphorus 14 mg
Riboflavin (B2) 0.1 mg Calcium 35 mg
Niacin (B3) 0.4 mg Iron 0.4 mg
Pantothenic acid (B5) 0.3 mg Magnesium 17 mg
Vitamin B6 0.1 mg Manganese 0.1 mg
Folate (B9) 6 mcg Potassium 232 mg
Vitamin C 2 mg Sodium 1 mg
Vitamin K 4.7 mcg Zinc 0.2 mg
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 705

respiration during fruit development. The low sucrose content might also be caused
by high invertase activity during fruit ripening. In contrast, several fruit species are
characterized by the accumulation of both sucrose and fructose in ripe fruit. The
latter is formed from sorbitol, translocated from leaves to fruit as sucrose (Vemmos
et al., 2013).
The sugar composition can influence fruit sweetness. For example, fructose has
a higher relative sweetness than glucose. Therefore, the perception of sweetness of
different fig accessions can be linked to their fructose content. The values in fig are
comparable to those reported in some other fruits, such as apples and strawberries.
Total sugar contentvaries among different fig cultivars (Veberic & Mikulic-
Petkovsek, 2016).

1.2 Nutrition and Health Considerations of Sugar Substitutes

1.2.1 Glycemic Index (GI)

Potassium is known to regulate blood sugar in the body. Fig is high in potassium and
keeps a check on blood sugar levels. Thus, the fig is said to help in maintaining
diabetes. Low potassium levels cause less insulin production, which leads to a spike
in blood sugar. Potassium affects the pancreatic β-cells, which are involved in insu-
lin secretions. Researchers have also found the lowering effect of chlorogenic acid
present in fig on blood sugar level. Chlorogenic acid reduces the blood glucose
concentration by inhibiting the G-6 phase, the two main metabolic pathways respon-
sible for the release of glucose from the liver (Ahrens & Thompson, 2013). Relative
glycaemic response to bulk sweeteners and alternatives show in Table 31.2.

Table 31.2 Glycaemic and insulinaemic responses to bulk sweeteners and alternatives (O’Donnell
& Kearsley, 2012)
Sugar or alternative Relative glycaemic response Categorisation
Maltodextrin 91 High
Maltose 105 High
Trehalose 72 High
Sucrose (present in fig fruits) 68 Intermediate
Lactose 46 Low
Isomaltulose 32 Very low
Glucose (main sugar of fig) 100 High
Fructose (main sugar of fig) 19 Very low
Tagatose 3 Very low
Xylitol 12 Very low
Sorbitol (present in fig fruits) 9 Very low
Maltitol 45 Low
Isomalt 9 Very low
Lactitol 5 Very low
706 A. Sharifi et al.

While non-communicable diseases are set to overburden private and governmen-

tal health budgets, preventive methods must maintain health. Dietary change pro-
vides one such method and explicitly reducing the glycaemic response to the diet
via food selection, food modification and ingredient choices and development are
valuable objectives. There is now growing evidence from clinical data confirms the
potential for reduced severity of disease, and from epidemiological data for reduc-
ing the risk of developing a variety of non-communicable diseases. Incorporation of
the low-glycemic approach as one component of a ‘better diet’ is likely to remain
important. Strategies that combine reduced GI or GL reduced saturated fat and
reduced energy are likely to be most effective, and these attributes are found in
alternative sugars and sweeteners (O’Donnell & Kearsley, 2012). The glycemic
index of figs is 51, which falls under the low GI category.

1.2.2 Dental Health

Fig syrup as a natural sugar substitute and low-glycemic sweeteners should have
little or no effect on a person’s blood sugar levels. However, applying natural sweet-
eners like fig syrup as a natural sugar substitute in food products can improve
oral health.
Low-calorie sweeteners are non-cariogenic; however, their low bulk in food,
drink, and medical products are often compensated by the inclusion of potentially
cariogenic bulking agents. Manufacturers, regulators, and consumers must recog-
nize any potential dental health risk. Therefore, investigation of the dental proper-
ties of sweeteners and sugar alternatives needs to continue and should be encouraged
for all potentially favorable new sweetening and bulking agents, especially those
likely to be used in food, drink, and medicine products likely to be consumed
between meals (O’Donnell & Kearsley, 2012).

1.2.3 Calorie Control and Weight Management

Some people choose to limit their food energy intake by replacing high energy sugar
with other sweeteners having little or no food energy (sugar substitutes). This allows
them to eat the same foods they usually would while losing weight and avoiding
other problems associated with excessive calorie intake (Tandel, 2011).
Non-nutritive, intense sweeteners have a valuable role in the diet providing
sweetness without calories. Using intense sweeteners to replace sugar can reduce
energy in foods and beverages, potentially energy intake. However, this depends on
the nutritional composition of particular foods and drinks and the physiological,
psychological, and behavioral effects of consuming foods with sweeteners in con-
trast to those with sugar. This chapter will look at the scientific evidence behind
these complex issues (O’Donnell & Kearsley, 2012).
Overall, evidence suggests that foods formulated with non-nutritive intense
sweeteners, reduced-calorie bulk sweeteners and bulking agents can play an
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 707

interesting and valuable role in helping consumers to improve the energy balance of
their diets and as part of weight reducing diets. Contrary to some reports, the main
body of published data shows that these ingredients, when incorporated into foods
with lower caloric density and/or reduced glycaemic impact, can help consumers to
eat fewer calories overall. A balanced approach to weight loss and maintenance is
essential for long-term success (O’Donnell & Kearsley, 2012).

1.2.4 Diabetes Mellitus

People with diabetes have difficulty regulating their blood sugar levels. However,
limiting their sugar intake by substituting sugar with artificial sweeteners allows
them to enjoy a varied diet; some sugar substitutes release energy but are metabo-
lized more slowly, allowing blood sugar levels to remain more stable (Tandel, 2011).

1.2.5 Hypoglycemic Effect

Hyperglycemia is any disease characterized by excessive urine secretion. The most

a common form of hyperglycemia is diabetes mellitus, the world’s largest endocrine
disease (Selvin et al., 2011). Fig fruit has an amazing property of regulating blood
glucose levels. Fig consumption results in a reduction in total cholesterol and a
decline in total cholesterol to HDL cholesterol ratio, which ultimately leads to a
decrease in hyperglycemia (Hussain et al., 2021).

1.3 Dried Fig Fruits

Dried fig is one of the most produced fruit products in the world. It is used in many
forms: as whole fruit for consumers; or in industrial forms, such as paste, concen-
trate, powder, and chopped fruit. Dried fig consumption manifests a positive effect
on human health. This beneficial effect has been ascribed to many micronutrients
reserved in dry fruit. Dried figs are rich in fiber, trace minerals, polyphenols, pro-
teins, sugars, and volatile compounds that provide a pleasant characteristic aroma
and are thus a good choice for a healthy snack (Bekatorou et al., 2002).
In addition to the energy provided by the high sugar content, dried figs have a
high nutrient content compared with other dried fruits. It is considered an important
source of minerals (potassium and calcium) and vitamins (B3 and nicotinic acid).
Dried fig fruits have a statistically higher individual and total sugar content than
fresh fruit (Table 31.3) (Veberic & Mikulic-Petkovsek, 2016).
Gebhardt et al. (1982) reported 52.90% total sugar and 51.3% reducing sugar
content in the dried fig fruit (Gebhardt et al., 1982). Thonte and Patil (1988) reported
that sugar-treated dried fig fruits contained 41.50–59.00% sugars. They reported
that ethereal-treated and dried figs contained 32.50% reduced sugar, whereas
708 A. Sharifi et al.

Table 31.3 Levels of individual sugars in dried figs (Veberic & Mikulic-Petkovsek, 2016)

Sugar Whole dried fruit (mg/100 g DW)

Sucrose 249–740
Fructose 8253–19,557
Glucose 9690–21,588
Total sugars 18,518-41,885

control fruits had 33.71% (Thonta & Patil, 1988). Pawar et al. (1992) reported that
the non-reducing sugar content in dried Poona fig was 6.64–9.82% (Pawar
et al., 1992).
Dried figs have one of the highest concentrations of polyphenols, a highly effec-
tive antioxidant. Fig syrup is just as good as the fresh and dried fruits if the right
preparation process is followed. When you opt for fresh fruit, you may have to wait
until it is in season and contend with the short storage period. The dried fruits may
stay for longer even though they can occupy a large storage space if you want to
keep a regular supply.

1.4 Sugar Replacement in Food Product

Sugars and sweeteners have an essential role in the human diet, and choosing the
right ones in the right amounts can influence health. Knowledge will enable good
choices, and further research and understanding of the literature will confirm or
deny how good our choices are and where improvements are possible. The choice is
not simply the healthier or healthiest since the technological properties and eco-
nomics of sugars and sweeteners impact which of them can be used suitably in a
particular food. A wide range of potential influences on health is offered by sugars
and sweeteners when selected appropriately (O’Donnell & Kearsley, 2012).

1.4.1 Synthetic Sugar Replacement in a Food Product

(High-Potency Sweeteners)

Six high-intensity sweeteners are FDA-approved as food additives in the United

States: saccharin (Hampton, 2008), aspartame (Tandel, 2011), acesulfame potas-
sium (Ace-K) (Tandel, 2011), sucralose (Tandel, 2011), neotame (Food &
Administration, 2018), and advantame (Tandel, 2011) (Table 31.4). High-intensity
sweeteners are commonly used as sugar substitutes or sugar alternatives because
they are often sweeter than sugar but contribute only a few to no calories when
added to foods. High-intensity sweeteners, like all other ingredients added to food
in the United States, must be safe for consumption (Food & Administration, 2018).
Table 31.4 Synthetic sugar replacement in food products
Sweetener Structure Regulatory status MSICS ADI NTSPE to ADIa CV EBNCS Reference
Acesulfame Approved as a 200× 15 23 0 Sweet One®Sunett® 1, 8, 3
Potassium (Ace-K) sweetener and
flavor enhancer in
foods generally
(except in meat
and poultry)
21 CFR 172.800
Advantame Approved as a 20,000× 32.8 4920 0 1, 5, 7, 8
sweetener and
flavor enhancer in
foods generally
(except in meat
and poultry)
21 CFR 172.803
Aspartame Approved as a 200× 50 75 4 Nutrasweet®Equal®Sugar 1, 2, 3, 8
sweetener and Twin®
flavor enhancer in
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute

foods generally
21 CFR 172.804
Neotame Approved as a 7000– 0.3 23(sweetness 0 Newtame® 1, 6, 8
sweetener and 13,000× intensity at
flavor enhancer in 10,000 × sucrose)
foods generally
(except in meat
and poultry)
21 CFR 172.829

(continued)
709
Table 31.4 (continued)
710

Sweetener Structure Regulatory status MSICS ADI NTSPE to ADIa CV EBNCS Reference
Saccharin Approved as a 200–700× 15 45(sweetness 0 Sweet and Low® Sweet 1, 5, 2, 8
sweetener only in intensity at 400× Twin® Sweet’N Low®
certain special sucrose) Necta Sweet®
dietary foods and
as an additive used
for specific
technological
purposes
21 CFR 180.37
Sucralose Approved as a 600× 5 23 0 Splenda® 1, 4, 5, 8
sweetener in foods
generally
21 CFR 172.831
Cyclamates 30–60× 11 0 1, 8

MSICS Multiplier of Sweetness Intensity Compared to Sucrose, ADI Acceptable Daily Intake: milligrams per kilogram body weight per day (mg/kg bw/d),
NTSPE to ADI Number of Tabletop Sweetener Packets Equivalent to ADIa, CV Caloric value (Cal/g), EBNCS Examples of Brand Names Containing Sweetener
References:
1-Carocho et al. (2017), 2- Meena et al. (2012), 3-George et al. (2010), 4-Furlán and Campderrós (2017), 5-Reis et al. (2011), 6-Morais et al. (2014),
7-Kobayashi et al. (2015). 8-Food and Administration (2018)
a
Number of Tabletop Sweetener Packets a 60 kg (132 pound) person would need to consume to reach the ADI. Calculations assume a packet of high-intensity
sweetener is as sweet as two teaspoons of sugar
®
Registered trademark
A. Sharifi et al.
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 711

The safety of High-Potency Sweeteners is demonstrated by a substantial body of

evidence and continued review by independent regulatory agencies/committees,
including JECFA/Codex, FDA, and EFSA. These organizations have considered the
decades of positive and negative human and animal studies to draw their conclu-
sions (Ashwell et al., 2020). Table 31.4 summarizes the most important synthetic
sugar replacement in food products.

1.4.2 Natural High-Potency Sweeteners

Many high-potency sweeteners of diverse chemical structures are known to occur

naturally. Natural sweeteners show many similarities to their synthetic counterparts’
overall taste characteristics. They range in potencies from approximately 10 times
that of sucrose to 1000s of times the sweetening power of sucrose. Overall sweet
taste quality, measured by traditional methods that assess temporal characteristics
and others that quantify the intensity of side- and aftertastes also range across the
spectrum in much the same way as has been found with synthetic sweeteners. As a
consequence, many of the challenges inherent in formulating sugar-reduced foods
and beverages so that they will deliver taste experiences fully acceptable to consum-
ers are similar, irrespective of the origin of the potent sweetener (O’Donnell &
Kearsley, 2012).
Natural high-potency sweeteners contain Brazzein and Curculin, which are low-
calorie sweeteners and are stable to heat and pH variation (DuBois & Prakash,
2012; Kant, 2005). Erythritol is a bulk sweetener for cooking, confectionery, and
beverages. It is stable to light and temperature, has a clean taste similar to sugar, and
has a cooling effect when tasted in solids (DuBois & Prakash, 2012; Kurihara &
Nirasawa, 1994; Priya et al., 2011). Glycyrrhizic acid is a natural flavor and flavor
enhancer. Glycyrrhizic acid has technological properties: soluble in hot water, anti-
inflammatory and anti-viral; bitter and licorice-like off-tastes; delayed onset of
sweetness and sweetness linger (Grenby, 1991; Priya et al., 2011). Other natural
high-potency sweeteners include the following: Mabinlin, Miraculin, Monatin,
Monellin, Pentadin, Steviol glycosides, stevioside and rebaudioside A, and
Thaumatin (DuBois & Prakash, 2012; Gibbs et al., 1996; Grenby, 1991; Kant, 2005;
Khattab et al., 2017; Kurihara & Nirasawa, 1994; Kroger et al., 2006; Priya et al.,
2011; Rodrigues et al., 2016). Table 31.5 shows the Potency of natural sweeteners.
Polyols (sugar alcohols) are nutritive sweeteners obtained by the catalytic hydro-
genation of the oxo-group of natural sugars, i.e., by substituting an aldehyde or keto
group with hydroxyl. The sweetness of sugar alcohols (polyols) is shown in
Table 31.6. The sweetness of polyols is lower than sucrose. Therefore, polyols
might be used as a bulk sweetener. Food products’ desired sweetness and flavor are
achieved by combining polyols and non-nutritive, usually artificial, sweeteners.
Polyols are responsible for texture, preservation, filling, moisture capture, and cool-
ing effect in the mouth. Polyol sweetness, such as maltitol, is up to 90% of the
sucrose sweetness (Petković, 2019).
712 A. Sharifi et al.

1.4.3 Plants and Fruits as a Source of Natural Sugar Replacement

in Food Products

The production of sugar-free foods, in which a sweet taste is obtained by using non-
caloric intensive sweetening substances, is becoming increasingly popular. Natural
high-intensity sweeteners are presented according to the source of origin. The plant
parts from They were obtained, and their sweetness potency was also considered
(Table 31.7). The sweetening properties, advantages, and disadvantages resulting
from the possibility of natural high-intensity sweeteners used in the production of
food is described, including sweeteners already in use or whose commercialization
is in progress (Świąder et al., 2019).

1.5 Fig Syrup

Fig industrialization produces many co-products. These co-products come from

fresh figs discarded because of inadequate ripening or over-ripening, spoiled, size,
texture, or low quality as table fruit, but they are safe for human consumption. The
non-use of fig co-product constitutes a real economic loss since it is rich in nutrients
and bioactive compounds, which can be extracted and used as value-added materi-
als. In addition, figs and figs co-products are an excellent source of minerals, vita-
mins, and dietary fiber; they are fat and cholesterol-free and contain many amino
acids (Viuda-Martos et al., 2015).
In formulating healthy food products, the fig fruit is an excellent source. The fig
is rich in vitamins, minerals, and dietary fiber and contains many natural antioxidant
compounds, according to the USDA. Furthermore, figs’ high fructose and glucose
content make them suitable for formulating foods with natural sweeteners (Chang
et al., 2016). However, because of economic reasons, damaged and deformed figs
are more applicable in the industry as they can be converted to fig extract and fig
powder. So it is vital to select those extraction conditions that may not cause heat
damage to nutrients and inhibit temperature-dependent adverse reactions such as
the Maillard reaction (Razavi et al., 2010).
The main challenge of fig syrup extraction is to provide an appropriate gentle
process. Based on documented assays, there are few reports concerning the produc-
tion process of fig extract (Farahnaky et al., 2010). Fig extract, however, does not
have a long shelf life. Therefore, it needs to be concentrated (Slatnar et al., 2011).
Iran produced 98,990 tons of fig fruit in 2020. The Sabz cultivar belonging to the
Smyrna type figs is the most common exportable Iranian local fig, among other
varieties, and Fars province, notably Estahban district estimated to be the most
important producer of sabz fig in the country (Kalantari et al., 2018).
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 713

Table 31.5 The potency of natural sweeteners (Potency is expressed as times sweeter than
sucrose) (Carniel Beltrami et al., 2018)
Sweetener Potency* Sweetener Potency*
Brazzein 500– Mogrol glycosides, Luo Han Guo sweetener, and ~250 (1, 8, 10)
2000 mogroside V
Curculin 550– Monatin 2700 (1)
9000
Erythritol 0.6–0.7 Monellin 2500–3000 (8,
10)
Glycyrrhizic 30–110 Pentadin ~500 (4, 6, 8)
acid
Mabinlin 100–400 Steviol glycosides, stevioside and rebaudioside A 200–400 (1, 8)
Miraculin 400,000 Thaumatin 1600–9800 (1,
7)
Potency is expressed as times sweeter than sucrose
*

Table 31.6 The basic important physical and chemical parameters of different polyols (sugar
alcohols)
Sweetness
Polyol (2–5) Food application (1)
Xylitol 1.0 Jellies, chewing gums, coatings for gum, mint-flavor candies
Maltitol 0.9 Chocolate, spread, hard candies, chewing gums, coating for gums
Sorbitol 0.6 Chewing gums, tablets, candies, humectants, plasticizers, hard
candies, baked goods
Erythritol 0.6 Hard/soft candies, chocolate, beverages, bakery products, chewing
gums
Mannitol 0.6 Dusting power, chewing gums, effervescent products
Isomalt 0.5 Chewing gum, dusting powder
Lactitol 0.4 Candies, frozen desserts, jams and jellies, chocolate, dusting powder,
bulking agent, baked products
Sucrose 1.0
References: 1: Petković (2019). 2: O’Brien-Nabors (2016). 3: Wheeler and Pi-Sunyer (2008). 4:
Fitch and Keim (2012). 5: Grabitske and Slavin (2008)

1.5.1 Fig Syrup Production

Several steps are involved in the production of fig syrup. First, figs are soaked in
water before a special mixer chops them up. Then, the mixture is lightly steamed
while careful tending helps check its consistency and keep it from overcooking.
This process is repeated several times to ensure that it gets to the right thickness.
Finally, the mixture will be sieved off and packed in storage containers ready for
distribution. Typical syrup can last for about a month when stored in a cool, dry
place without refrigeration. Therefore, the syrup is mainly safe for most people and
can be added to different foods. Fig syrup, owing to the particular sweet and sour
taste, can be adopted either in cake production (as filler and topping) or as a sort of
‘barbecue sauce’; therefore, it is important to guarantee the rheological characteris-
tics suitable for different applications (Gabriele et al., 2010).
714 A. Sharifi et al.

Table 31.7 Characterization of plants and high-intensity sweeteners obtained from them (Świąder
et al., 2019)
FDA/EFSA
Plant Sweetness sweetener
Plant species parts Sweeteners potency* Compound status
Capparis masaikai Fruit Mabinlin I-1 10–400 Protein Not approved
lev. Seeds Mabinlin II
Mabinlin III
Mabinlin IV
Curculigo latifolia Fruit Curculin 500** Protein Not approved
Dryand (Lemba) Curculin 430–2070
Dioscoreophyllum Fruit Monellin 3000 Protein Not approved
cumminsii (Stapf)
Diels
(SerendipityBerry)
Pentadiplandra Fruit Pentadin brazzein 500–2000 Protein Not approved
brazzeana Baill.
Synsepalum Fruit Miraculin 400,000** Protein Not approved
dulcificum Daniell
(miracle fruit)
Thaumatococcus Fruit Thaumatin I 1600– Protein GRAS flavor
daniellii (Benn.) thaumatin II 3000 enhancer/E957
Benth (Katemfe) thaumatin a sweetener,
thaumatin b flavor
thaumatin c enhancer
Glycyrrhiza glabra Root Glycyrrhizin 50 Terpenoid GRAS flavor
Linn (Licorice) (triterpenoid Enhancer/not
glycoside) approved EU
Siraitia grosvenorii Fruit Siamenoside I 500 Terpenoid GRAS/not
Swingle (Luo Han 11-oxomogroside V (triterpenoid approved EU
Guo, monk fruit) Mogroside V glycoside)
mogroside IVa
mogroside IVe
Stevia rebaudiana Leaf DulcosideA 30 Terpenoid GRAS steviol
Bertoni (stevia) RebaudiosideA 200 (diterpenoid glycosides/E
RebaudiosideB 150 glycoside) 960
RebaudiosideC 30 steviol
RebaudiosideD 221 glycosides
RebaudiosideE 174 sweetener
RebaudiosideF 200
RebaudiosideM 250
Rubusoside 114
Stevioside 210
Steviolbioside 90
Sclerochiton Root Monatin2R,4R 3000 Indole Not approved
ilicifolius A. Meeuse Monatin 2S,4S 1200
(Arruva)
relative to sucrose
*

modifying sour taste into sweet taste

**
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 715

Kalantari et al. (2018) optimize extraction conditions and fig syrup production at
three different temperatures (5, 25, and 40 °C), Fig to water ratios of (1:1.0, 1:1.5,
and 1:2.0 w/w) and storage times of 12, 24, 48, and 72 h. The optimum extraction
condition of fig extract was provided by a temperature of 5 °C, a fig to water ratio
of 1:1.5 w/w, and an extraction time of 72 h, which produced fig extract with
14% ± 0.5°Brix. The extract was then passed through a sieve (with a mesh size of
400 μ) so that large suspended solids were removed from the liquid (Kalantari et al.,
2018). Therefore, fig extract with 14% °Brix needs to be concentrated to increase its
shelf life. Table 31.8 shows the physicochemical properties of fig extract.
During the production of fig syrup, fig fruits were ground and then mixed with
boiling water for the time requested for the solution to reach a standard concentra-
tion (14% °Brix). After filtration, the extracted solution is concentrated by evapora-
tion, reducing the water content to the desired level. Under usual industrial
conditions, during concentration, the atmospheric pressure is maintained at 1 atm,
and the temperature of the boiling solution increases above 100 °C because of boil-
ing point increase with syrup concentration. These process conditions are used for
simplicity and low cost, even though either flexibility or control of final syrup prop-
erties are penalized (Gabriele et al., 2010).
Process innovation is oriented toward correcting weak points by modifying the
process conditions to either a more selective extraction of components from fruits or
better process control, improving flexibility. The efficiency of solid-liquid extraction
can be improved by reducing raw material dimensions: owing to the increase in sur-
face/volume ratio, the contact between solid and solvent is more efficient, and some
components can be more easily extracted. High-temperature conditions are com-
monly used because they can increase mass transfer rate and thermodynamic solubil-
ity limits of some components, yielding to faster processes (Gabriele et al., 2010).

Table 31.8 Physico-chemical properties of fig (Ficus carica L.cv. Sabz) extract (Kalantari
et al., 2018)
Properties Fig extract
Total soluble solids (1 g/100 g of the fig extract) 14.0 ± 0.3
Total solids (1 g/100 g of the fig extract) 15.34 ± 0.04
Protein 0.28 ± 0.04
Fat 0.09 ± 0.03
Total sugar (1 g/100 g of the fig extract) 6.81 ± 0.11
Glucose 3.7 ± 0.2
Fructose 2.8 ± 0.1
Acidity (based on the concentration of citric acid) 0.30 ± 0.02
pH 4.88 ± 0.12
Ash (1 g/100 g of the fig extract) 0.67 ± 0.09
L* 50.66 ± 2.06
a* −1.33 ± 0.76
b* 20.33 ± 0.11
L*(Lightness)
a* (Red/Green Value)
b* (Blue/Yellow Value)
716 A. Sharifi et al.

However, high temperatures cause thermal degradation of some fruit compo-

nents, mainly polysaccharides, leading to highly reactive intermediates that can
form colored polymers (Coca et al., 2004). As a consequence, nutritional properties,
rheological properties and appearance are affected by drastic thermal conditions,
and milder processes should be used to preserve some product characteristics even
though a longer operating time could be necessary. The main components extracted
from fruit are simple sugars (mainly fructose and sucrose) and complex polysac-
charides. In addition, it is known that long-chain branched molecules build up a 3-D
network into the material as a result of the gelation process and chain-chain interac-
tions; therefore, consistency is mainly related to the amount of extracted polysac-
charides like starch, a mixture of amylose and amylopectin, and pectin, a complex
polymer containing D-galacturonic acid units (Gabriele et al., 2010).
When considering the latter, owing to the chemical behavior of its carboxylic
groups, it is possible to improve the efficiency of extraction by changing the pH
value and adopting a slightly alkaline solvent. Consequently, a more or less struc-
tured syrup could be obtained by varying the extraction pH. Therefore, the tradi-
tional process was modified from these general considerations, and the following
changes were introduced (Fig. 31.1).
1. Addition of sodium bicarbonate and ammonium bicarbonate as alkaline agents.
2. Addition of a preliminary fruit grinding to increase the extraction efficiency.
3. Adoption of low temperature to reduce thermal degradation reactions.
4. Under-vacuum concentration to reduce operating temperature and oxygen avail-
ability for oxidation processes. Adopting the proposed changes, a bench-scale
process was set up as a scale-down of traditional industrial systems, and different
syrup samples were prepared to evaluate the effects of each adopted modifica-
tion (Gabriele et al., 2010).
Gabriele et al. (2010) investigated the effects of selected operating parameters on
the fig syrup rheology. Process conditions have been changed, either adding alkaline
salts, as pH modifiers to improve pectin extraction efficiency, or decreasing opera-
tive temperature, reducing thermal damage of the material. It has been found that
alkaline salt addition yields structured products because of the improved efficiency

Fig. 31.1 Process flow sheet for figs syrup production. (Gabriele et al., 2010)
31 Fig (Ficus carica) Syrup as a Natural Sugar Substitute 717

of pectin extraction and the enhancement of the gelation phenomena. In addition,

mild conditions during either extraction or concentration lead to a more structured
syrup, whereas more drastic operations bring liquid-like products (Gabriele
et al., 2010).
In addition, it is interesting to notice that it is possible to adopt the process condi-
tions to produce an industrial-standard syrup always having the same rheological
properties, even when raw materials properties can enormously change because of
external factors such as weather conditions, fruit cultivar, and harvesting period.
This ‘rheology-aided’ process analysis allows for directly choosing the proper con-
ditions, reducing significant time- and money-consuming ‘trial and error’ proce-
dures focused on standardization of high-quality product production (Gabriele
et al., 2010).

1.5.2 Powder Production of Fig or Fig Syrup

The ready-to-eat foods in the food market are mostly powdered; therefore, an appro-
priate method to dry figs will increase their usage in various products. As a result,
powder technology is gaining importance in the food industry and among food pro-
ducers (Koç & Ertekin, 2016). Spray drying, tray drying, freeze-drying, foam mat
drying, and microwave drying are the most used drying methods to produce pow-
dered products. However, after drying (except for spray drying), the obtained prod-
uct is scraped and milled to obtain powder (Varhan et al., 2019).
Fig extract does not have a long shelf life. Therefore, it must be concentrated or
converted to powder with longer-lasting storage characteristics. Drying is the major
food processing operation that can enhance the shelf life (Slatnar et al., 2011).
Spray drying has a short-timed process and operational conditions that can be
maintained. It is broadly used to produce high-quality powders regarding water
activity, color, flavor, and nutritional values from various foods and fruit and vege-
table extracts. For example, spray drying of fig extract can be used to produce
healthy foods because it is considered some encapsulation (Kalantari et al., 2018).
The spray drying process is very effective in preparing any easy-to-use and long-
lasting powder that, after reconstitution, is more or less similar to the original juice,
commonly known to be directly related to drying conditions (Encina et al., 2016).
Sharifi et al. (2015), Zareifard et al. (2012), Jamdar et al. (2021), and Shahidi et al.
(2020) have published some valuable reports regarding the operating conditions and
qualitative characteristics of spray-dried juice/extract powders (Sharifi et al., 2015;
Zareifard et al., 2012; Jamdar et al., 2021; Shahidi et al., 2020).
The foam mat drying method consists of converting liquid or semiliquid foods
into stable foams via intense agitation and air and using foaming agents and stabiliz-
ers. As liquid foams are metastable, stabilizing agents are needed, usually surfactant
molecules. Stabilizers delay the several mechanisms of foam aging, such as drain-
age, coalescence, and coarsening. Some foam stabilizers include xanthan gum, pro-
pylene glycol alginate, methylcellulose, Arabic gum, and maltodextrin. In foam-mat
drying, drying usually refers to dehydration of the thin foam layer by hot air. Besides
718 A. Sharifi et al.

foam mat hot-air drying, other drying methods, including freeze-drying, vacuum-
drying, infrared-drying, and microwave drying, have also been reported (Hajiaghaei
& Sharifi, 2022).
Varhan et al. (2019) optimized foam mat-dried fig powder formulation. The fig
foam was dried through hot air and microwave separately. Their results showed that
the microwave drying method was more efficient than the hot air in time. Besides,
the higher foam thickness provided more hydroxymethylfurfural content in the fig
powder. Therefore, the drying method and conditions were important to produce a
high-quality foam mat-dried fig powder (Varhan et al., 2019).
Khapre et al. (2015) produced fig (Ficus carica L.) fruits powder, and the fig
powder was utilized in a value-added product like burfi (Indian cookie). Fig powder
incorporated burfi was nutritionally rich in terms of fiber (3.7%), potassium (0.46%),
and protein (13.1%). The prepared product was low-cost compared to similar mar-
ket products (Khapre et al., 2015). Khapre et al. (2011) also produced toffee with fig
fruits powder (Khapre et al., 2011). Yeganehzad et al. (2020) investigated the pos-
sibility of developing a fruit snack formulation based on dried fig powder and
chocolate-coated (Yeganehzad et al., 2020).

2 Conclusions

Consumer interest in natural high-potency sweeteners has grown in recentyears,

followed by concerns about the use of artificial additives in foods. Knowledge of
potent sweeteners in plants is not new; there have been literature references to
examples dating back to the mid-nineteenth century. However, most references to
potent natural sweeteners also note overall taste characteristics incompatible with
successful commercialization. Fig syrup is a rare example of a super-food that also
tastes great. It has been appreciated for centuries in the Mediterranean and the
Middle East and is now appreciated worldwide. The health benefits of figs in gen-
eral and of fig syrup, in particular, have been known for thousands of years. Figs are
far more nutritious than some commonly used natural sweeteners like dates. Fig
syrup can be used as the potential substitute for sugar in various food products such
as fruit juices, biscuits, cakes, muffins, desserts, and confectionery.

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Chapter 32
Fig (Ficus carica) Shelf Life

Elham Taghavi, Akram Sharifi, Navideh Anarjan, and Mohd Nizam Lani

Abbreviations

1-MCP 1-Methylcyclopropene
APE Aqueous soy polyphenolic antimicrobial extract
APX Ascorbate peroxidase
CA Controlled atmosphere
CDB Cuello Dama Blanco
CDN Cuello Dama Negro
CP Cold plasma
DNA Deoxyribonucleic acid
EW Electrolyzed water
FDA Food and Drug Administration
GRAS Generally Recognized As Safe
Gy Gray
MAP Modified atmosphere packaging
OA Ozonated air
OTR Oxygen transmission rate
RH Relative humidity
ROS Reactive oxygen species
TSS Total soluble solids

E. Taghavi (*) · M. N. Lani

Faculty of Fisheries and Food Science, Universiti Malaysia Terengganu,
Kuala Nerus, Malaysia
e-mail: [email protected]
A. Sharifi
Department of Food Science and Technology, Faculty of Industrial and Mechanical
Engineering, Qazvin Branch, Islamic Azad University, Qazvin, Iran
N. Anarjan
Department of Engineering, Tabriz Branch, Islamic Azad University, Tabriz, Iran

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 723
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_32
724 E. Taghavi et al.

WVTR Water vapor transmission rate

1 Shelf Life

Shelf life is the time for which a stored item remains usable (International Food
Information Service, 2007), or in other words, shelf life is the length of time food
could be kept before it deteriorates (Lombarda, 2006). Fruits are unique among food
products in terms of shelf life since they remain as living tissues until consumed or
processed. All living tissues respire, and the consequences are quite profound for
these products’ shelf-life and storage stability. Slowing respiration can slow senes-
cence and thus prolong shelf life; however, some respiration must continue, or the
products will rapidly senesce and die. Furthermore, each fruit is unique in genetic
makeup (species, cultivar, clone, etc.), its stage of development (maturation, stage
of ripening, etc.), and the pre-and post-harvest conditions it has experienced. These
facts make a shelf-life prediction of fresh produce particularly difficult compared to
products with more uniform composition and stability (Aked, 2001).

2 Fig Shelf Life

Fresh figs should be plump and relatively soft but free of bruises (Murdock, 2002).
Fig fruit is highly delicate at harvest and quite perishable after harvest and during
handling and cold storage (Byeon & Lee, 2020). It should be refrigerated for 7 days
(Murdock, 2002). After harvest, the quality of fig fruit is relatively unstable, and the
fruit’s fresh weight loss and respiration rate are sharply increased during cold stor-
age, regardless of the cultivar (Byeon & Lee, 2020). This fruit is generally catego-
rized as climacteric fruit as they show a sharp increase in respiration rate and
ethylene production at the end of phase II of the double sigmoid growth curve
(Rosianski et al., 2016). When fig fruit mature, their dimensions (diameter and
length) abruptly increase, contributing to the rupturing of the ostiole-end region and
peel tissues (Kong et al., 2013). These fig fruit physiological disorders are referred
to as ostiole-end splitting and peel side cracking, respectively (Villalobos et al.,
2016), and they deteriorate fruit quality after harvest and during cold storage by
favoring conditions that induce fruit decay (Byeon & Lee, 2020; Villalobos
et al., 2016).

3 Factors Influencing the Shelf Life of Fig

Various factors influence the shelf life of the fig. These factors include fig character-
istics, microbiological spoilage, chemical deterioration, physical deterioration,
transportation and distribution, temperature, storage atmosphere, insects and
32 Fig (Ficus carica) Shelf Life 725

packages, and moisture. Figure 32.1 shows the factors that influence the shelf
life of fig.

3.1 Fig Characteristics

Appearance is the key factor for consumers in making purchases of fresh produce.
Vital components of visual quality include color and color uniformity, glossiness,
absence of defects in shape or skin finish, and freedom from disease. Textural prop-
erties and flavor and aroma of fig are other criteria that define fig as fruit (Aked,
2001). Fig shape is usually obovoid, turbinate, or pear-shaped. Its size may range
from 2.5 to 10 cm in length. Fig color varies depending on the variety; some of the
typical colors are yellow, green-yellow, copper color, red and purple. The skin is
tender and thin, and the syconium wall is fleshy and either pale yellow, amber, light
pink, red, or purple (Crisosto et al., 2011). Generally, the fig is a pleasantly sweet

Fig. 32.1 Factors influencing the shelf life of fig

726 E. Taghavi et al.

fruit that consists of soft flesh pursed around a large number of tiny edible seeds
(Murdock, 2002). Fig texture is a very important parameter for consumer accep-
tance. a texture that is too soft may be rejected, and it can increase the degree of
mechanical injury (Bahar & Lichter, 2018). In addition, the quality of the fig could
affect its shelf life of the fig.

3.2 Microbiological Spoilage

Fresh figs are very sensitive to microbial spoilage, even in cold storage conditions.
Thus, fresh figs are high perishable products during postharvest with microbiologi-
cal decay that induces an unpleasant taste and smell due to rot, and suitable conser-
vation methods must be applied (Paolucci et al., 2020). The growth and proliferation
of microorganisms cause the main postharvest losses in fig fruits. The predominant
microflora of fresh fig is mainly composed of moulds such as Botrytis cinerea,
Monilinia laxa, Alternaria alternata, Fusarium moniliforme, Rhizopus stolonifer,
Aspergillus niger, Cladosporium herbarum, and Phytophora palmivora, being most
of them responsible of fruit decay (María del Carmen Villalobos et al., 2017).
Among the main diseases reported in fig fruits, we find endosepsis caused by
Fusarium spp. or smut caused in dried fruit by Aspergillus niger, Alternaria rot
(caused by Alternaria alternata or other Alternaria spp). Additionally, these moulds
can be pathogenic or toxigenic due to their mycotoxins production ability while
growing on fruits (Cantín et al., 2011; María del Carmen Villalobos et al., 2017),
even during its cold storage. Likewise, yeast such as Hanseniaspora spp., and
Torulopsia spp. have been reported as the main yeast species in fruits (María del
Carmen Villalobos et al., 2017; Tournas & Katsoudas, 2005). Specifically, species
belonging to the genus Hanseniaspora, Saccharomyces, Pichia or even bacteria
such as Bacillus may cause souring or fermentation (Cantín et al., 2011; María del
Carmen Villalobos et al., 2017).
Other microbial groups such as mesophilic aerobic, lactic acid bacteria (LAB),
Staphylococcus spp., Enterobacteriaceae spp., Pseudomonas spp., and Acetobacter
spp. have been described to a lesser extent as part of the microbial population in ripe
fruits and vegetables, which can also be responsible for fruit decay and health haz-
ard (Badosa et al., 2008; María del Carmen Villalobos et al., 2017).

3.3 Chemical Deterioration

Chemical spoilage may be via non-microbial enzymatic action, oxidation, or non-

enzymatic browning. Physical spoilage includes water loss, increased moisture in
dry food, freezer burn, and recrystallization of frozen foods. However, there are
occasions when the cause and/or manifestation of spoilage is a combination of these
32 Fig (Ficus carica) Shelf Life 727

different types of hazards, e.g., physical changes to the product or its container
caused by chemical reactions or microbial growth and metabolism (Blackburn, 2006).

3.4 Physical Deterioration

Fresh figs are very perishable (Crisosto et al., 2011), sensitive to physical damage,
and highly susceptible to postharvest decay infections (Crisosto et al., 2011; Venditti
et al., 2005). Fresh fig quality (taste and aroma) increases with ripening; however,
its sensitivity to damage also increases, reducing its market life (Crisosto et al.,
2011). Figs have concise storage and shelf life. This is mainly due to their fast ripen-
ing and high susceptibility to decay (Crisosto et al., 2011; Venditti et al., 2005),
which results from their easily damaged epidermis and high sugar content. The fruit
epidermis is very sensitive to pressure and knocks, which can cause fig bruising,
splitting and injuries, favoring pathogen infections. Sensitivity to physical damage
is affected by the fig cultivar and the maturity stage at harvest (Crisosto et al., 2011).

3.5 Transportation, Distribution, Temperature, Storage

Atmosphere, Insects and Package

• Transportation & distribution

The effective distribution of perishable commodities like fresh figs is of utmost
importance when transporting. Currently, mismanagement of premium fruit and
vegetables leads to a considerable amount of capital loss, such that logistics service
providers apply refrigerated trucks to deliver them to minimize the food spoilage
rate during transportation (Tsang et al., 2018). Moreover, fruits and vegetables
retain their physiological functions, such as respiration and transpiration, after har-
vesting and remain fresh only as long as their normal metabolism continues (Prasad
et al., 2018). So, attention to these issues is quite a concern during transportation
and distribution.
• Temperature
Low cold storage is one of the strategies used to maintain quality and extend the
shelf life of fruit (Mditshwa et al., 2020). Postharvest temperatures are critical for
fresh figs (Crisosto et al., 2011). The number of studies dealing with postharvest
technologies for fresh figs is somewhat limited, but it is generally accepted that stor-
age temperature should be low (0–2 °C) and relative humidity (RH) should be high
(Bahar & Lichter, 2018; Gözlekçi et al., 2005). For instance, in California, ‘Black
Mission’ and ‘Calimyrna’ figs, two different types of figs stored in optimum condi-
tions (−1 to 0 °C), have an expected postharvest life of 1–2 weeks in air and
3–4 weeks in a controlled atmosphere (Crisosto & Kader, 2007).
728 E. Taghavi et al.

• Storage atmosphere
Storage in a controlled atmosphere (CA) with 5–10% oxygen and 15–20% carbon
dioxide is beneficial for reducing decay, maintaining firmness, and reducing respira-
tion and ethylene production rates. Reducing respiration and ethylene production
rates will lengthen fresh fruit life (Crisosto & Kader, 2007; Crisosto et al., 2011).
However, extended storage in CA could lose characteristic flavor (Crisosto et al.,
2011). Bahar and Lichter (2018) studied CA as a potential tool to extend the shelf
life of Ottomanit figs. They found that storing Ottomanit figs under CA conditions
of 5 kPa O2 and 5 kPa CO2 maintains fruit quality better than the ambient atmo-
sphere or a CA with higher CO2 levels. This study improved the quality concerning
firmness, integrity, and decay control.
• Insects
Souring or fermentation spoilage of the figs occurs when the fruit is still on the tree.
The agents that cause souring are yeasts and bacteria, which insects transport.
Therefore, souring symptoms are only noticeable when fruits are ripe and have the
ostiole wide open. The ostiole aperture is necessary for the presence of this disease
since the ostiole is the way of entrance for the vector insects. Therefore to control
this disease, it is necessary to eradicate the vector insects (Crisosto et al., 2011).
• Package
The package protects its contents from outside environmental effects, such as water,
water vapor, gases, odors, microorganisms, dust, shocks, vibrations, compressive
forces, etc., and protects the product’s environment. For many food products, the
protection afforded by the package is an essential part of the preservation process.
In general, once the integrity of the package is breached, the product is no longer
preserved (Robertson, 2016).

3.6 Moisture

Water is the most abundant constituent of food. It is a key determinant of food stor-
age and has many roles in processing (Hssaini et al., 2020; Kong & Singh, 2016). In
macroscale chemistry, the water impact on food reactions and quality is more
important than any chemical constituent (Rahman, 2007). The importance of water
function in biological products goes far beyond its quantitative aspect. Water con-
tributes to the texture and appearance of vegetables and plays an essential role in the
occurrence of biochemical reactions and microbial growth (Berk, 2018). Controlling
the food moisture content during processing is an ancient preservation technique
that is still widely practiced today (Desa et al., 2019). This is achieved by either
moisture removal or binding to ensure long food stability to microbial and chemical
deterioration (Kong & Singh, 2016). During post-harvest, fresh fruits and vegeta-
bles continue to undergo changes leading to their senescence and deterioration, such
32 Fig (Ficus carica) Shelf Life 729

as biochemical reactions and microbial growth. These two factors are the most com-
mon limitation on food shelf life. Hence, several preservation processes aim to
achieve the stability of foods by maintaining the moisture content under the levels
required for microbial development and biochemical reactions. This stability mainly
depends on the relationship between the food equilibrium moisture content and its
corresponding water activity at a constant temperature (Hssaini et al., 2020; Peng
et al., 2007). Figs are an important source of essential dietary nutrients such as
fibers, carbohydrates, minerals, and vitamins (Hssaini et al., 2019; Hssaini et al.,
2020). Since the moisture content of fresh figs is generally greater than 80% (dry
basis), they are classified as highly perishable commodities (Desa et al., 2019;
Hssaini et al., 2020). Fig water loss can be reduced with early cold storage. However,
longer periods of delayed cooling tended to produce a lower percentage of quality
fruit during the storage life and shelf life of figs (Crisosto et al., 2011).

4 Techniques on Shelf Life Extension of the Fig

Various practical technology can reduce the fig losses and shelf-life extension of
figs. These include common or traditional techniques (drying, using chemical pre-
servatives such as 1-methylcy-clopropene, calcium chloride, chlorine dioxide, and
packaging) and novel techniques (edible coating, irradiation, ozone, cold plasma,
electrolyzed water, and antimicrobial). Figure 32.2 illustrates the Techniques for
shelf life extension of fig.

4.1 Common Techniques on Shelf Life Extension of Fig

4.1.1 Drying

As mentioned earlier, the moisture content of fresh figs is generally greater than
80% (dry basis); they are classified as highly perishable commodities (Desa et al.,
2019; Hssaini et al., 2020). Moreover, fresh and dried figs can adsorb or desorbs
water during storage depending on the environmental conditions, which, if not con-
trolled, this water loss and gain can negatively affect their quality. Therefore, their
valorization depends on maintaining the moisture content under the levels required
for its spoilage or deterioration. Therefore, nowadays, an important interest is given
to drying and monitoring the moisture content changes during and after this process
as one of the most suitable fruit preservation methods (Hssaini et al., 2020).
Moreover, drying can extend the fig shelf life remarkably. More information can be
found on fig drying technologies in this book.
730 E. Taghavi et al.

Fig. 32.2 Techniques on shelf life extension of fig

4.1.2 Chemical Treatments

Chemical treatments have been shown to affect weight loss during the storage and
shelf life (Mditshwa et al., 2020). Therefore, some studies investigate the effects of
chemical substances applied on figs to improve their postharvest conservation, such
as 1-methylcy-clopropene (1-MCP) and calcium chloride.
• 1-Methylcyclopropene (C4H6)
1-MCP is an unsaturated cyclic olefin that acts as a competitive ethylene antagonist,
i.e., it blocks ethylene receptors and can be used to control ethylene production,
respiration rate, and softening and extends the shelf-life of a wide range of fruits
(Brasil Dias Tofanelli et al., 2018). However, 1-MCP applications on fruits may not
always have similar results on the postharvest quality since its effect has been shown
to vary according to the climacteric fruit species, cultivar, maturation and ripening
stages of the fruit, and 1-MCP application forms (Brasil Dias Tofanelli et al., 2018;
Junhua Zhang et al., 2017). 1-MCP was patented in 1996, followed by rapid regis-
tration and commercialization because of its non-toxic mode of action, effective-
ness at low concentrations, and easy application as a gas. It has also been used
extensively as a research tool to study the effects of ethylene on a range of climac-
teric and nonclimacteric fruits, vegetables, and flowers (Li et al., 2016; Jing Zhang
et al., 2020). The impact of 1-MCP on the physiological and biochemical processes
related to fruit ripening and consequent effects on quality during storage and shelf-
life periods could vary greatly among different types of fruits, storage durations,
and storage conditions. The majority of commercial use of 1-MCP is on apples,
with less use on other products, in part because it is harder to delay rather than
totally inhibit ripening of fruits such as avocado, banana, pear, and tomato (Jing
Zhang et al., 2020).
There are few studies on the postharvest behavior of figs treated with
1-MCP. However, Gözlekçi et al. (2005) revealed that 10 μg/L of 1-MCP slowed
down the softening of Bardakci figs and retained fruit firmness during a period of
32 Fig (Ficus carica) Shelf Life 731

15 days of storage in styrofoam trays wrapped with commercial cling film at 0 °C

and 90–92% RH. Also, D’Aquino et al. (2003) treated Bianca figs with 400 μg/L of
1-MCP at 20 °C for 24 h and concluded that 1-MCP treatment of the figs and stor-
age at 15 °C for 7 days might improve their storability.
In the other study, Brasil Dias Tofanelli et al. (2018) investigate the effects of
different concentrations of 1-MCP on postharvest quality of Roxo-de-Valinhos
fresh ripe figs. Their research was conducted by applying 4 various concentrations
of 1-MCP (0 as control, 5, 10, and 20 μg/L) on figs, after which the fruits were
evaluated at five different storage times (0, 4, 8, 12, and 16 days). The result showed
that the1-MCP treatment improved the quality (firmness and acidity) of ripe Roxo-
de-Valinhos figs during storage. In this regard, 20 μg/L of 1-MCP provided the best
fruit firmness at day 12 after harvest. Higher acidity was observed in the figs with-
out a 1-MCP treatment at 8 days of storage and with 20 μg/L of 1-MCP after 12 days
of storage. Moreover, 1-MCP did not affect fruit weight loss. Overall, the figs
treated with 1-MCP showed promising and rising values of firmness, acidity, and
total soluble solids (TSS) at the highest dose. The TSS concentration tended to
increase at the higher doses of 1-MCP.
• Calcium chloride (CaCl2)
The postharvest calcium treatment effectively reduced physiological diseases
and delayed senescence of fruits (Gao et al., 2020; Zhang et al., 2019). In addition,
calcium treatment effectively delays ripening and maintains the quality of many
fruits (Gao et al., 2020). The postharvest calcium treatment is a safe and non-toxic
alternative to chemical treatment for fruit storage and has become an important
focus for research in fruit preservation (Gao et al., 2020; Mishra et al., 2018). It
maintains cell shape, cell membrane integrity, tissue hardness, and lipid metabolism
and prolongs fruits‘storage period. However, the possible mechanism of calcium
regulating fruit ripening is still unclear (Gao et al., 2020).
Irfan et al. (2013) studied the response evaluation of fig fruit to dip treatments
with calcium and sodium salts to improve the fruit texture and retain initial fruit
quality attributes during storage and extend storage life. They found that CaCl2
extends the keeping quality of fig fruit during storage and shelf-life. CaCl2 in 4%
concentration was notable in retention of fruit color, texture, and increased accumu-
lation of ascorbic acid, compared to untreated control figs. Besides, pretreatment
with CaCl2 in 4% concentration was most effective in growing both mesophilic
aerobic bacteria and yeast and molds at low temperature (1 ± 0.5 °C, 95–98% RH)
storage; it caused further delays in ripening and senescence of figs. Therefore 4%
CaCl2 was beneficial in prolonging the postharvest life twofold.
• Chlorine dioxide (ClO2)
Chlorine dioxide is a potent oxidizing agent and has a broad antimicrobial activ-
ity. It has about 2.5 times the oxidation capacity of chlorine. In addition, it does not
react with nitrogen-containing compounds or ammonia to form dangerous chlora-
mine compounds. Therefore, the US Food and Drug Administration (FDA) (1998)
732 E. Taghavi et al.

has allowed the use of aqueous chlorine dioxide in washing fruit and vegetables
(Karabulut et al., 2009).
Karabulut et al. (2009) examined the possibility of using chlorine dioxide by
fogging to control postharvest diseases fig. In their research, the fruit was fogged
with various concentrations of chlorine dioxide in a cold storage unit for 60 min at
room temperature. Treated fruit was stored either in the air or in modified atmo-
sphere bags for 7 d at 1 °C followed by 2 days shelf-life at 20 °C. Fogging at
300–1000 μL/L significantly reduced the natural incidence of decay, most of which
was gray mold. Modified atmosphere packaging did not improve the efficacy of
fogging in reducing decay incidence. The epiphytic population on the fruit surface
was similarly reduced by chlorine dioxide fogging. All treatments significantly
reduced total microorganisms, fungal and bacterial populations in fig fruit. In addi-
tion, microorganisms in the storage atmosphere were significantly reduced. None of
the treatments affected the visual quality and taste of fruit.

4.1.3 Packaging

• Modified atmosphere packaging (MAP)

MAP is a technique used for prolonging the shelf life of fresh or minimally pro-
cessed food by changing the composition of the air surrounding the food in the
package (Mangaraj & Goswami, 2009). In other words, MAP is the imposition of a
gas atmosphere, typically containing an inert gas, such as nitrogen, combined with
an antimicrobial active gas, such as carbon dioxide, upon a packaged food product
to extend its shelf life (Spencer, 2005). In MAP, the modification of the atmosphere
inside the package is achieved by the natural interplay between the product’s respi-
ration and the permeation of gases through the packaging. Also, the gas composi-
tion is modified initially and changes dynamically depending on the respiration rate
of the food product and the permeability of the film surrounding the food product
(Mangaraj & Goswami, 2009).
Furthermore, modifying the atmosphere within the package can be established
either passively by commodity or intentionally via active packaging or a combina-
tion of the both (Mahajan et al., 2007; Mangaraj & Goswami, 2009). In the case of
active MAP, an atmosphere is established by pulling a slight vacuum and replacing
the atmosphere of the package with the desired gas mixture of O2, CO2, and N2. On
the other hand, in the case of passive MAP, the modification of the atmosphere
inside the package is achieved by natural interplay between two processes, the res-
piration of products and the transfer of the gases through the packaging that leads to
an atmosphere richer in CO2 and poorer in O2 and it depends on the characteristics
of the commodity and packaging film (Mangaraj & Goswami, 2009).
Fresh figs have high respiratory intensity (64 mg CO2 /kg.h) at ambient tempera-
ture (25 °C) (Liao et al., 2016), which causes their daily water loss rate to exceed
5% (Ye, 2014). Hence, modified atmosphere packaging (MAP) is one of the most
appropriate ways to maintain a fig’s quality. Studies by Ma et al. (2019) have
32 Fig (Ficus carica) Shelf Life 733

designed two types of MAP: DMAP and TMAP. DMAP consists of O (O film is a
modified PE film by styrene butadiene styrene block polymer) and W film (W film
is a modified PE film by nano vermiculite), whereas TMAP consists of B, M, and W
film. B and M film refer to modified O film (B film) whose OTR is 4.44 × 105 mL/
m2 and 1-MCP releasing the film, respectively. O and B film has a high oxygen
transmission rate (OTR) and low water vapor transmission rate (WVTR). On the
other hand, W film has high WVTR and low OTR. Therefore, low oxygen, high
carbon dioxide, and high humidity atmosphere were formed in DMAP and
TMAP. Their studies showed that DMAP and TMAP could prolong shelf life of figs
from 2 days to 4 days and up to 6 days at 25 °C. In addition, figs with DMAP and
TMAP maintained a lower loss of water, firmness, decay rate, TA content, vitamin
C content, and high sensory quality during storage.
Also, the use of packaging under a modified atmosphere has been reported to
reduce the respiration rate, with the benefit of delaying senescence by reducing
metabolic activity and microbial proliferation (Valero et al., 2008). These effects of
MAP have been previously reported by Maria del Carmen Villalobos et al. (2014)
for fresh breba fruits and figs. (2016), who reported that the use of microperforated
films to create passive modified atmospheres was effective in reducing the incidence
of decay caused by fungal growth and in retaining quality characteristics. Thus,
atmospheres with low levels of O2 and high levels of CO2 inhibit the growth of most
aerobic microorganisms due to the well-known antimicrobial activity of CO2 at high
concentrations (María del Carmen Villalobos et al., 2017; Serradilla et al., 2013).
Martinez-Damian et al. (2020) report on MAP‘s effect on nutraceutical quality
and overall appearance of figs stored at 1 °C. The combined use of packaging and
cold in their investigation allowed them to observe fig with less weight loss and
firmness, maintaining the citric acid and Vitamin C values without significant
changes. Except for the last evaluation period, no statistical variation was found in
the content of total anthocyanins, total phenols, and antioxidant capacity. Moreover,
it was also possible to observe a significant impact on the reduction of respiration
and ethylene production, which could be corroborated by the conservation of sen-
sory aspects of the fig (texture, color, appearance, marketing level, aroma, and fla-
vor). The generation of passive atmospheres (packaging) and cold (1 °C) are
valuable tools in maintaining the nutraceutical quality and appearance of fig fruits,
aspects highly appreciated by consumers and marketers of this fruit.

4.2 Novel Techniques on Shelf Life Extension of Fig

4.2.1 Edible Coating

Edible coatings are mainly used to increase food products appearance and extend
the shelf life of the fruits (Salehi, 2020). By definition, edible coatings are natural
food-safe and ecologically friendly substitutes applied to reduce water transfer, gas-
eous exchange, and oxidation of fresh produce (Tesfay et al., 2017). A sustainable
734 E. Taghavi et al.

approach to extending the shelf-life of figs can be constituted by applying an edible

coating able to maintain the quality of the fruit during storage (Paolucci et al.,
2020). Also, edible coatings have a high potential to carry active ingredients such as
anti-browning agents, colorants, flavors, nutrients, and antimicrobial compounds
that can extend product shelf life and reduce the risk of pathogen growth on the food
surface (Galus et al., 2020; Vieira et al., 2021).
Paolucci et al. (2020) studied the active edible polysaccharide-based coating to
preserve fresh figs. They compared fresh figs in a commercial preservation system
with the figs preserved in an edible coating (Alginic acid/Agar, 70/30) and an active
edible coating to preserve their quality characteristics. The coating efficacy was
enhanced with the addition of pomegranate peel extract at two different concentra-
tions (0.25% and 0.5%). Including a component with high antioxidant activity in an
edible coating proved to be an excellent method for preserving the quality of this
highly perishable fruit, such as fig. In the other research, Vieira et al. (2021) inves-
tigated the enhancement of postharvest quality and shelf life of fresh figs via com-
posite coatings of chitosan and alginate emulsions with olive oil during storage at
4 °C. Their results revealed that coatings effectively delayed postharvest ripening
indicators (i.e., respiration rate, mass loss, softening, total soluble solids/titratable
acidity ratio) and fungal decay. The results foresee a fruit’s shelf life between 14 and
19 days under refrigeration at 4 °C that may be followed up to 2 days at ambient
temperature, higher than that estimated for uncoated fruits (less than 14 days at 4 °C
plus to 2 days at ambient temperature).
Adiletta et al. (2019) studied the chitosan coating to preserve the physicochemi-
cal (weight loss, soluble solid content, and titratable acidity) and nutraceutical traits
(total polyphenol, anthocyanina, flavonoid, ascorbic acid content, and antioxidant
capacity) in fresh fig (Troiana). They compared fruits treated with 1% chitosan and
1% ascorbic acid coating and subjected to refrigerated storage (4 °C) for 9 days.
Their results have shown that alginate-chitosan bilayer coating had a significant
effect on weight loss of fresh fig since weight loss for coated only 5–7% is much
lower than uncoated fig, which 18%. Their finding also demonstrated that chitosan
coating delayed the decrease in ascorbic acid content in fresh fig fruits during cold
storage. Moreover, chitosan-coated figs also maintained a higher antioxidant capac-
ity than uncoated ones during 9 days of cold storage. The catalase activity had
decreased significantly in uncoated fig while showing a stable trend toward storage
in coated fig. Ascorbate peroxidase (APX) activity had increased significantly in
chitosan-coated fruits compared to the uncoated fig during cold storage. Their study
highlighted the beneficial effects of the antioxidant system that allowed to delay
oxidative damage during postharvest treatment.

4.2.2 Irradiation

The application of irradiation can improve safety and extend the shelf of fruit like
fig. Irradiation can use for various purposes such as: eliminating foodborne illness
effectively, destroying or inactivating organisms that cause spoilage and
32 Fig (Ficus carica) Shelf Life 735

consequently extending the shelf life of foods, controlling the insects by decreasing
the need for other pest-control practices that may harm the fruit, and delaying in the
ripening of fruit to increase longevity. There are three sources of radiation approved
for use in foods by the FDA: Gamma rays, X-rays, and electron beams (or e-beam)
(Administration, 2016).
In a study related to fig irradiation, Silva et al. (2009) investigated the physico-
chemical characteristics changes caused by γ-irradiation in pre-ready green figs to
increase the useful shelf life of fig products. In their study, the samples were pro-
cessed (washing, making hygienic, cooking for 15 minutes, and cooling), wrapped
in a plastic sack, and stored at 8 °C in an OBD camera for 7 days. Later samples
were irradiated with doses of 0 (control), 1.0, and 2.0 kGy, under a dose of
0.601 kGy/h, in a Gammacell-220 irradiator and stored for 24 h to 8 °C in
OBD. These results confirmed that the irradiation did not significantly alter the
physicochemical properties (i.e., pH, soluble solids content, color peel, color pulp,
texture, chlorophyll A, chlorophyll B, and total carotenoids) of pre-ready green fig,
with the exception for texture in all doses. In this regard, more research on irradia-
tion needs to be undertaken for fig shelf life.

4.2.3 Ozone

Ozone (O3) is a powerful sanitizer that is Generally Recognized as Safe (GRAS). In

2001, ozone, in gas and aqueous phases, was approved by the US Food and Drug
Administration (FDA) as an antimicrobial additive for direct contact with foods. In
the handling process, ozone can be applied in two forms. First, gaseous ozone is
added continuously or intermittently to the storage atmosphere of the harvested
product. Aqueous ozone is added immediately after the harvest or during the wash-
ing treatment. Extrinsic and intrinsic factors influence the efficacy of ozone.
Extrinsic factors include pH, organic matter, pressure and temperature (in water
quality), air relative humidity (in air quality), concentration, and treatment time
application method (in ozone treatment); while intrinsic factors include the type of
fruit and vegetable, weight, characteristics of the product surface, and surface area
(in a food product), the activity of water (aw), characteristics of microbial strains,
physical state of bacterial strains, natural microflora, artificially inoculated microor-
ganisms, and population size (in microbial load) (Sarron et al., 2021).
Some researchers believe that fresh figs cannot be washed as their skin can be
easily damaged during the washing process, and water may induce the germination
of fungal spores on the surface of the fruit (Afsah-Hejri et al., 2021; Karabulut et al.,
2009). In this context, Afsah-Hejri et al. (2021) investigated the effect of ozonated
air (OA) on weight loss, volume reduction (shriveling), and skin firmness on fresh
figs and monitored the changes in the epidermis of OA-treated fruits within 14 days
of storage at 4 °C and 65% RH. For this purpose, Kadota and Black Mission figs
were treated with OA (15 ppm for 3 h), and their aging parameters were evaluated.
Their study showed that OA treatment of fresh figs could be used to minimize the
volume and weight loss and improve the overall quality of fresh figs.
736 E. Taghavi et al.

In the other study, Zorlugenç et al. (2008) studied the effectiveness of gaseous
ozone and ozonated water on microbial flora and aflatoxin B1 content of dried figs.
Dried figs were exposed to13.8 mg/L of ozone gas and 1.7 mg/L ozonated water for
7.5, 15, and 30 min. During the ozonated water process, E. coli, coliform, yeast, and
molds were completely inactivated, whereas approximately an 88% reduction was
observed in aerobic mesophilic bacteria count at the end of the 30 min. The E. coli
and molds population was destroyed in gaseous ozone treatment. However, a sub-
stantial reduction in aerobic mesophilic bacteria, coliform, and yeast counts was
determined. In addition, Aspergillus flavus and Aspergillus parasiticus, which cause
aflatoxin formation, were isolated from non-ozonated dried figs. Due to the inacti-
vation of all molds in dried figs, aflatoxin formation potential was decreased after
the ozonation process. Overall, the experimental results of their research indicated
that the application of gaseous ozone and ozonated water had a significant effect on
the degradation of aflatoxin B1 in contaminated dried figs.

4.2.4 Cold Plasma (CP)

Cold plasma (CP), also known as non-thermal discharge plasma, is an emerging

non-thermal processing technique that ensures food quality and safety (Waghmare,
2021). CP is an emerging green process technology in today’s industrialized era
since it is eco-friendly and chemical-free (Chen et al., 2019; Waghmare, 2021).
Plasma is generally characterized as “the fourth state of matter”. Plasma consists of
complete or partly ionized gases with a range of charged and neutral reactive spe-
cies that drive its chemistry (Varilla et al., 2020). CP is defined as a source of high-
energy electrons at ambient temperature and pressure that interact in an open or
controlled environment (Varilla et al., 2020; Whitehead, 2016). As a novel non-
thermal technology, CP can be applied to keep the freshness of the fruit. Therefore,
it was practical to extend the shelf life of fruit by inhibiting its softening. Also, the
low temperature and active particles during CP processing play a significant role in
reducing food nutrient loss and biomacromolecules modification. Therefore many
studies about applications of CP for food processing have been reported for decon-
tamination, enzyme inactivation, starch modification, toxin degradation, food
allergy, and food packaging (Pan et al., 2021).
The action mood of CP for food decontamination is back to the principal active
components identified for the antimicrobial activity of CP systems are reactive oxy-
gen species (ROS) (Varilla et al., 2020; Whitehead, 2016). CP also produces hydro-
gen peroxide and nitrogen species with antimicrobial properties (Maisch et al.,
2012; Varilla et al., 2020). Furthermore, the effect of plasma treatment on microbial
cells is mainly due to the plasma ions and cell interactions. The reactive species in
plasma have been widely associated with the direct oxidative effects on the outer
surface of microbial cells. The effect of plasma is highly dependent on the presence
of water. The highest effect was observed in moist organisms compared to the low-
est in the dry organism. The potential application of plasma in inactivation is based
32 Fig (Ficus carica) Shelf Life 737

on the fact that reactive plasma species damage the deoxyribonucleic acid (DNA) in
the chromosomes (Thirumdas et al., 2015).
Burana (2021) reported the effect of plasma treatments and low temperature on
some fresh fig fruit‘s physical and physiology. In this investigation, various times of
plasma treatment, including 0 (control, untreated), 2, 5, and 10 hours during storage
at 4 °C, were considered. It was found that plasma treatments reduced the metabo-
lism of fresh figs, including weight loss, respiration, and ethylene production during
storage. Also, the decrease of firmness was reduced by plasma treatment, especially
during 2 h plasma treatment. Moreover, plasma treatments considerably inhibited
fungal incidents that prolong the storage life of fresh figs under storage at
4 °C. Unfortunately, though, these treatments did not affect the color change in this
fruit. The findings recommend that plasma treatment combined with storage at low
temperature has commercial potential and helps reduce postharvest decay, main-
taining the quality and prolonging the storage life of fresh fig fruit.
In the other research, Abbaszadeh et al. (2018) studied the application of CP
technology in the quality preservation of fresh fig fruit. In this study, dielectric bar-
rier discharge plasma was applied to fruits in two steps. Durations of the first treat-
ments were 1 and 5 min. Based on these results, main experiments were conducted
for 30, 90, and 180 s, and CP was applied to packed and unpacked fig samples.
Subjective measurements showed significant shelf-life improvement for the treated
figs compared to the control samples, while objective quality attributes were not
altered except for pH and a* index which were not undesired. Therefore, direct
application of plasma for 90 s and in-package treatment for 30 s were suggested for
further investigation in their research. Furthermore, pulsed plasma processing of
packed figs for 30 s is recommended considering the practical aspects. Overall they
conclude that atmospheric CP has the potential to be applied to fresh fig fruits to
prolong their shelf life.

4.2.5 Electrolyzed Water (EW)

EW refers to the acidic or alkaline solution produced by the electrolysis of various

neutral salts in an electrolytic cell (Zhang et al., 2021). In this technology, by using
tap water and salt, the system generates two-stream simultaneously, one is alkaline,
and the other is acidic, with the characteristics of cleaning and sanitizing solutions,
respectively. Researchers evaluated the effectiveness of these solutions as sanitizing
and cleaning agents (Yamaner et al., 2016). The EW is a sustainable and green con-
cept and has several advantages over traditional cleaning systems, including cost-
effectiveness, ease of application, effective disinfection, on-the-spot production,
and safety for human beings and the environment (Rahman et al., 2016). The main
applications of EW in improving the quality of postharvest fruits and vegetables
include controlling the microorganisms in postharvest, suppressing browning,
delaying senescence, maintaining bioactive compounds, alleviating chilling injury,
and removing pesticide residues from fresh fruits and vegetables. Therefore, EW
738 E. Taghavi et al.

treatment seems to be a promising method for preserving postharvest fruits and

vegetables (Zhang et al., 2021).
There is no study on the application of EW in postharvest figs so far. However,
Yamaner et al. (2016) studied the efficacy of neutralized EW and mild heat against
foodborne pathogens isolated from fig. In their study, spores of Aspergillus flavus
and Penicillium expansum were isolated from the surface of fig fruits. In addition,
Escherichia coli and Bacillus cereus, known on figs’ surfaces, were also evaluated.
Spores and vegetative cells of fungi and bacterium were exposed to five different
concentrations of neutralized EW (100, 75, 25, 5, and 1%) at three different tem-
peratures (22, 50, and 70 °C) for 1, 3, and 5 min. According to the results, at 22 °C,
1% neutralized EW exposure for 1 min effectively decreased the number of E. coli
and B. cereus vegetative cells by approximately 8.5 log cfu/mL and 6.3 log cfu/mL,
respectively. At 50 °C, 5% neutralized EW exposure for 1 min significantly reduced
the P. expansum and A. flavus spore numbers by 7 log cfu/mL and 5.54 log cfu/mL,
respectively. With mild temperature (22–50 °C), low chlorine neutralized EW
(9.22 mg/L - 33.85 mg/L available chlorine concentrations) showed a strong anti-
bacterial and antifungal activity against foodborne pathogens. They conclude that
neutralized EW can be used widely as a sanitizer in fig enterprises instead of high-
cost chlorine-based disinfectants.

4.2.6 Antimicrobials

Only a few investigations and limited studies have been focused on using natural
antimicrobial agents from plants and microorganisms to extend the shelf life of fig.
Fig crops are distinguished by their perishability and searching for antimicrobial
agents that can extend the fruits‘shelf life are very challenging. Besides mold,
mesophilic aerobic, lactic acid bacteria (LAB), Staphylococcus spp.,
Enterobacteriaceae, Pseudomonas spp., and Acetobacter spp. are also responsible
for fruit decay of fig; however, they have been described in a little attention (Badosa
et al., 2008).
The use of aqueous extracts obtained from the roots, leaves, or seeds of plants
has been proven to extend the shelf life of fruits. Maria del Carmen Villalobos et al.
(2016) have studied the soybean meal extract was applied by dipping two fig culti-
vars; Cuello Dama Blanco (CDB) and Cuello Dama Negro (CDN), in combination
with microperforated M50 film (1/50 mm; ø = 100 μm) as well as with macroperfo-
rated film in order to evaluate the gas composition, weight loss, percentage of fruits
with disorders, microbial counts, physicochemical parameters and sensory quality
during cold storage for 21 days. Their studies showed the synergistic effects when
combining soybean meal extract with MAP can reduce mold and yeast levels,
reduce disorders by fungal growth and maintain the quality parameters of fig culti-
vars. For the CDN cultivar, cold storage could be extended to 21 days regarding
physicochemical quality and control of postharvest decay, while the use of M50
film without natural compounds application allowed the shelf life up to 14–17 days
for the CDN cultivar and 7 days for CDB cultivar.
32 Fig (Ficus carica) Shelf Life 739

Other technologies such as biocontrol agents and natural compounds such as

essential oils or aqueous phenolic extracts have been used in food preservation due
to their antimicrobial properties (María del Carmen Villalobos et al., 2017; Isman,
2000; Janisiewicz et al., 2001). In addition, these compounds inhibit the mycelial
growth of molds and spore germination, affecting the cellular metabolism of the
pathogens (María del Carmen Villalobos et al., 2017; Regnier et al., 2010; Maria
Serrano et al., 2005; Tzortzakis, 2007a, b). Moreover, the combination of both tech-
nologies renders them more effective in prolonging the postharvest life and the
safety of the horticultural produce (María Serrano et al., 2008; Villalobos et al., 2015).

5 Future Trend

Various practical technology can reduce the fig losses and shelf-life extension of
figs. These include common or traditional techniques (drying, using chemical pre-
servatives such as 1-methylcy-clopropene, calcium chloride, chlorine dioxide, and
packaging) and novel techniques (edible coating, irradiation, ozone, cold plasma,
electrolyzed water, and antimicrobial agents). Future work could evaluate the pos-
sibility of increasing the figs shelf life using traditional and novel techniques.

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Chapter 33
Use of Proteolytic Activity of Ficus carica
in Milk Coagulation

Hasitha Priyashantha, C. S. Ranadheera, Tharindu R. L. Senadheera,

H. T. M. Hettiarachchi, Shishanthi Jayarathna, and Janak K. Vidanarachchi

Abbreviations

Cys Cysteine
MMTS Methylmethanethiol sulfonate
mPEG Monomethoxypolyethylene glycol
ORAC Oxygen radical absorbance capacity
PLCPs Papain like cysteine proteases

1 Introduction

Proteases are hydrolytic enzymes that could hydrolysis the peptide bonds in pro-
teins into smaller polypeptides or amino acids, and at present, approximately 60%
of the commercialized enzymes are proteases (Mazorra-Manzano et al., 2018). This

H. Priyashantha (*) · S. Jayarathna

Department of Molecular Sciences, Swedish University of Agricultural Sciences,
Uppsala, Sweden
e-mail: [email protected]; [email protected]
C. S. Ranadheera
School of Agriculture & Food, Faculty of Veterinary & Agricultural Sciences,
The University of Melbourne, Parkville, VIC, Australia
e-mail: [email protected]
T. R. L. Senadheera
Department of Biochemistry, Memorial University of Newfoundland, St. John’s, NL, Canada
e-mail: [email protected]
H. T. M. Hettiarachchi · J. K. Vidanarachchi
Department of Animal Science, Faculty of Agriculture, University Peradeniya,
Peradeniya, Sri Lanka
e-mail: [email protected]; [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 745
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_33
746 H. Priyashantha et al.

group of enzymes plays a vital role in many industries, including food, pharmaceu-
tical, detergent, leather, etc., and therefore, has received significant commercial
interest. Even though the various synthetic, animal, plant, and microbial-derived
proteases are abundant, much focus has been recently directed towards plant-derived
proteases as they possess broad substrate specificities and activities over a wide
range of temperature and pH (Kumari et al., 2010; González-Rábade et al., 2011).
Plant proteases play an essential role in plant growth, reproduction, digestion,
seed germination, defense mechanism, apoptosis, etc. Hence, contribute to every
aspect of the plant life-cycle, from mobilizing storage proteins during seed germina-
tion to cell death (Schaller, 2004). The quantity, quality, and type of plant proteases
vary from one species to another and/ or within the same plant itself (Ben Amira
et al., 2017). This greater diversity among thousands of plants has resulted in vari-
ous plant proteases having different functionalities. Nearly all parts of plants, e.g.,
fruits, flowers, latex, leaves, stems, seeds, and roots, contain proteases and have
been extracted using different methods (Shah et al., 2014). MEROPS, the peptide
database (http://merops.sanger.ac.uk), has classified proteases into seven families
according to their active sites. Among them, plant proteases belong to five families:
(I) Serine, (II) Cysteine (Cys), (III) Aspartic, (IV) Metallo, and (V) Threonine
(Rawlings et al., 2006).
Papain, bromelain, and ficin derived from Carica papaya, Ananas comosus, and
Ficus spp., respectively, are the most widely used commercialized plant endo-
proteases that belong to the Cys-family (Devaraj et al., 2008a, b). In the food indus-
try, they are mainly used in meat tenderization, dairy processing, brewing, beverage
production, flavored-protein hydrolysates, emulsifiers, etc. (Devaraj et al., 2008a, b;
González-Rábade et al., 2011). Among those applications, plant proteases such as
milk coagulants have become a great interest in the dairy industry.
Milk coagulation is the most crucial primary step in the cheese-making process,
with much economic interest among cheese producers. Generally, milk coagulation
achieves using calf rennet, a mixture of chymosin and pepsin (Anusha et al., 2014).
However, the increasing world population, economic growth, and changes in eating
behaviours have considerably raised the need to expand cheese production capaci-
ties and efficiencies. Nevertheless, the supply of rennet is not sufficient to meet the
current demand due to the low availability of ruminant abomasa (Roseiro et al.,
2003; Jacob et al., 2011; Ben Amira et al., 2017). Furthermore, the emerging vegan
market, due to the ban of recombinant products in some countries (e.g., The
Netherlands, France, Germany, etc.), animal welfare concerns, the trend of vegetari-
anism, religious taboos, and health concerns coupled with the limited supply of
rennet have encouraged the use of alternative plant proteases in milk coagulation
(Roseiro et al., 2003; Aider, 2021).
Moreover, easy access, availability, low cost, and simple extraction and purifica-
tion procedures have attracted more attention to discovering novel plant proteases as
cheap alternatives for the calf rennet. As a result, numerous plant proteases have
been studied worldwide for their milk clotting properties. Among those various
plant proteases, ficins (also known as ficain) derived from Ficus carica L. have
gained much attention as milk coagulants due to their well-characterized structures,
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 747

functions, and properties diversified and precise usage in milk coagulation at the
industrial level. Therefore, this chapter examines the use of ficins derived from
F. carica L. in milk coagulation by paying detailed attention to its properties, extrac-
tion, mechanisms, and characteristics.

2 An Overview of Use of Plant Proteases in Milk Coagulation

Milk coagulation is essential in the cheese-making process and has long been stud-
ied due to its commercial and research interests. Proteolytic enzymes (e.g., chymo-
sin) destabilize the casein micelles sterically stabilized in their native form in milk
due to the “hairy” κ-casein surface layer. Chymosin (EC 3.4.23.4) cleaves at the
specific peptide bond of Phe105-Met106 of protruding κ-casein protein by releasing
caseinomacropeptide (κ-casein peptide 106–169) to the aqueous phase of the milk
(serum) and thereby weakening the steric and electrostatic stabilization of micelles
(Fox et al., 2017). Milk coagulation properties directly influence the properties of
the resulting cheese.
Rennet is a mixture of chymosin and pepsin (EC 3.4.23.1) discovered long before
from the abomasum of suckling calves (Roseiro et al., 2003). High specificity
towards κ-casein and low calf rennet proteolytic activity made it the most suitable
enzyme for cheese production (Anusha et al., 2014). However, the high price and
the limited supply of calf rennet and ethical, religious, and dietary protests led to
discovering possible alternatives to coagulate milk. An alternative milk coagulant
should have similar biochemical properties, such as greater specificity towards
κ-casein, high milk clotting index, proteolytic activity at the cheese making pH, and
temperature to calf rennet (Liburdi et al., 2019). Moreover, it should have sufficient
thermostability to ensure proper whey drainage without residual coagulant activity.
Even though microbial and recombinant chymosins are available, plant proteases as
milk coagulants have become a growing interest in the cheese industry due to their
great diversity and clean-label environment in usage.
Plant proteases have been used as milk coagulants in artisanal cheese production
over a long period in the Mediterranean, West African, and Southern European
countries (Liburdi et al., 2019). Almost all parts of plants contain proteases that can
clot the milk under appropriate conditions (Roseiro et al., 2003; Shah et al., 2014).
Many plant proteases which show milk clotting properties belong to aspartic prote-
ases (Table 33.1). However, some cysteine and serine proteases have also shown
milk clotting properties. These proteases have been extracted from different parts of
plants (Table 33.1), such as latex, leaves, flowers, stems, fruits, and seeds (Roseiro
et al., 2003; Shah et al., 2014).
Although numerous proteases are in the findings, only a few are on the industrial
applications, such as papain, bromelain, cardosins, and ficins, as their structures,
functionalities, and required conditions for various food processing operations have
been thoroughly evaluated. Among them, ficins from F. carica have gained much
748 H. Priyashantha et al.

Table 33.1 Types and sources of plant proteases having milk clotting properties
Type of
protease Source Tissue References
Aspartic Balanites aegyptiaca Fruits Beka et al. (2014)
Bromelia hieronymi Fruits Bruno et al. (2010)
Centaurea Germinated Salvador et al. (2006)
calcitrapa seeds
Cirisum vulgare Flowers Lufrano et al. (2012)
Citrus aurantium Flowers Tripathi et al. (2011) and Mazorra-
manzano et al. (2018)
Cynara humilis Flowers Esteves et al. (2003)
Cynara scolymus Flowers Chazarra et al. (2007)
Cynara cardunculus Flowers Roseiro et al. (2003), Silva et al. (2003),
Gomes et al. (2018)
Moringa oleifera Flowers Pontual et al. (2012)
Onopordum Flowers Brutti et al. (2012)
acanthium
Oryza sativa Seeds Asakura et al. (1997)
Solanum Fruits Guiama (2010) and Chávez-garay et al.
elaeagnifolium (2016)
Solanum tuberosum Leaves, tubers Tito et al. (2020)
Streblus asper Stems Salehi et al. (2017)
Withania coagulans Fruits Cavalli et al. (2008)
Cysteine Actinidia chinensis Fruits Puglisi et al. (2014)
Albizia lebbeck Seeds Egito et al. (2007)
Asclepias Latex Liggieri et al. (2009)
curassavica
Bromelia pinguin Fruits
Calotropis gigantea Latex, stems Anusha et al. (2014)
Calotropis procera Latex, leaves Aworh and Muller (1987) and Abebe and
Emire (2020)
Euphorbia nivulia Latex Badgujar and Mahajan (2014)
F. carica sylvestris Latex Faccia et al. (2012)
Ficus racemosa Latex Devaraj et al. (2008a, b)
Helianthus annus Seeds Egito et al. (2007) and Jain et al. (2020)
Sideroxylon Latex
obtusifolium
Zingiber officinale Rhizome Huang et al. (2011)
Serine Crinum asiaticum Latex Singh et al. (2010)
Cucumis melo Fruits Shah et al. (2014)
Cucurbita moschata Seeds Kumar and Sasmal (2020)
Ficus religiosa Latex Kumari et al. (2010)
Lactuca sativa Leaves Lo Piero et al. (2002)
Solanum dubium Seeds Ahmed et al. (2009)
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 749

attention since their milk clotting properties have been characterized and assessed
thoroughly.

3 Chemistry, Composition, and Properties of Proteases

in F. carica

3.1 Ficins (EC 3.4.22.3); the Major Proteolytic Enzyme

in F. carica

F. carica L., generally known as “Common Fig” is a tree with deeply lobed (three
or five lobes) leaves (Fig. 33.1a), which belong to the family Moraceae. Fig is a
latex-producing fruit crop and consists of numerous varieties with considerable
genetic diversity (Badgujar et al., 2014; Barolo et al., 2014). Although figs are
native to Southwest Asia and the Mediterranean region, fig cultivations are wide-
spread globally (Badgujar et al., 2014). Figs are among the earliest cultivated fruits,
are highly nutritious, and possess cultural, medicinal, and industrial values; thus
have a high demand (Devaraj et al., 2008a, b; Badgujar et al., 2014). Apart from the
fruit (Fig. 33.1b, c), fig latex is the essential component in Ficus spp., possessing a
wide range of industrial usage due to its inherent proteolytic activity.
Generally, plant latices are complex emulsions rich in proteases, alkaloids,
starches, sugars, oils, tannins, resins, gums, etc. (Kumari et al., 2010). For example,
fig latex consisted of 70% water, 20% gum, 9% ficins, and 1% other dry materials
(Zare et al., 2013). The reason for latex having such a high number of proteases and
their exact roles is still unclear. However, findings suggested that they are involved

Fig. 33.1 F. carica, also known as Common Fig plant’s healthy leaves (a), immature fruits (b),
and matured/ripened fruits (c) (Photo: Hasitha Priyashantha)
750 H. Priyashantha et al.

in defense mechanisms by protecting fruits against plant pathogens (Kumari et al.,

2010; Azarkan et al., 2011; Haesaerts et al., 2015). In addition to ficin (EC:3.4.22.3),
which is the main protease in latex of fig (Barolo et al., 2014), various other second-
ary metabolites, i.e., cis-1,4-polyisoprene, alkaloids, terpenoids, tannins, sterols are
also found in fig latex (Milošević et al., 2020). Most of these compounds are toxic
to pathogens, including insets and fungi, and therefore, act as defensive chemicals
in fig fruit (Zare et al., 2013).
The majority of proteases found within plant latices belong to the cysteine and
serine proteases, and they encompass several proteases such as papain, bromelain,
ficin, calotropins, etc., in industrial applications. Ten proteolytic components named
A, B, C,…, J from the lattices of different F. carica varieties have been identified
(Sgarbieri et al., 1964). Among them, the Cys endo-proteases, commonly termed
ficins (as illustrated in Fig. 33.2), which are closely related to the papain family
(family C1, clan CA), are the most exploited industrial proteolytic component
(Haesaerts et al., 2015). The proteases in latex of F. carica are available in multiple
forms of proteolytic enzymes, including ficin, lysozyme, class-II chitinases, and a
thaumatin-like protein (Baeyens-Volant et al., 2015). Proteases or proteolytic
enzymes play a fundamental role in hydrolyzing or cleaving peptide bonds of pro-
tein, which is used as a beneficial enzymatic activity in food processing, e.g., milk
coagulation, meat tenderization, dough conditioning agent, bioactive peptide pro-
duction, etc. (Aider, 2021).

Fig. 33.2 Latex of fig contains 9% of ficin (EC:3.4.22.3), the major proteolytic enzyme. Its chem-
ical structure (C9H14N4O3) (a) and 3D structure (b) are visualized using 4YYU in the protein
databank; https://www.rcsb.org/structure/4YYU. (3D image is made with PyMOL v1.8.6.0
(Schrödinger LLC). Photo: Hasitha Priyashantha)
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 751

Purified ficin is a unique single polypeptide chain with a molecular mass of

23.1 kDa (Devaraj et al., 2009). Moreover, most papain-like cysteine proteases
(PLCPs) are polypeptides with small molecular masses ranging from 20 to 30 kDa
(Rawlings et al., 2016). Nevertheless, it shares some common properties like sub-
strate specificity, transpeptidase reactions, activation by reducing agent, and ester-
ase activity similar to papain’s (Hamed et al., 2020). Therefore, ficin is also known
as one of the main enzymes in the group of PLCPs that includes bromelain, calpain,
cathepsin B, and chymopapain (Aider, 2021). Ficin consists of 201 amino acid resi-
dues (Aider, 2021). The amino acid sequence present in the active site residues of
ficin resembles the corresponding sequence of the papain (Milošević et al., 2019).
Ficin is a nonspecific sulfhydryl protease classified as a cysteine endopeptidase
enzyme (Zhai et al., 2021). Compositional analysis of purified ficin further indicates
the presence of three disulfide bonds and a single free cysteine residue at the active
site (Devaraj et al., 2008a, b; Rawlings et al., 2016). The active site of the ficin spe-
cifically consists of cysteine (Cys-25) and histidine (His-159) (Aider, 2021).
Several ficin isoforms from F. galabrata were isolated and purified in the early
1940s, and their structures have been thoroughly evaluated. Multiple ficin isoforms
from F. carica have also been isolated; however, some of these isoforms’ functions,
properties, and structures are yet to be discovered (Zare et al., 2016). Moreover, the
number and the relative amounts of ficin types available within different F. carica
cultivars and varieties also differed (Sgarbieri et al., 1964; Zare et al., 2016). To the
current knowledge, F. carica contains 6 ficin isoforms, namely Ficin A, B, C, D1,
D2 (Azarkan et al., 2011), and E (Baeyens-Volant et al., 2015). They all are non-
glycosylated molecules. Among them, Ficin A is the most abundantly available ficin
isoform. The recently discovered ficin E co-eluted with ficin A has a different
N-terminal sequence (LPETMDWRSK) and electrophoretic behavior compared to
other isoforms (Baeyens-Volant et al., 2015). All the other five isoforms have simi-
lar molecular mass and N-terminal ends; however, their crystallization structures
differ considerably. Ficin isoforms B and C share a common crystal formation, indi-
cating a close sequence and a structural similarity, while ficin A and D showed some
distant formations (Haesaerts et al., 2015). The protein data bank codes for ficin A,
B, C, and D were 4YYQ, 4YYR, 4YYV, and 4YYW, respectively (Milošević et al.,
2020). Recent investigations on ficin S revealed that it possesses a different isoelec-
tric point and a separate sugar content compared to other isoforms (Morellon-
Sterling et al., 2020). However, limited studies have been conducted to date to
characterize the physicochemical, kinetic, three-dimensional structures, and other
functional properties of different ficin isoforms (Aider, 2021). Ficin is mainly free
form, and ficin cocktail applications are becoming popular as an industrial biocata-
lyst (Morellon-Sterling et al., 2020).
Generally, ficins from F. carica have a wide specificity range toward essential
and neutral amino acids such as Gly, Val, Leu, Ala, Ser, Asn, Arg, His, and they
hydrolyze Tyr, Phe, and Val peptide bonds in proteins (Yang et al., 2017; Aider,
2021). Ficin has been recognized as sulfhydryl protease and contains two sulfhydryl
groups. Only one group directly reacts with the catalytic site, while the other group
occasionally shows its activity (Devaraj et al., 2008a, b; Zare et al., 2013). Moreover,
752 H. Priyashantha et al.

it is necessary to have -SH, -NH3+, and -COO− reactive groups to have catalytic
action of ficin (Zare et al., 2013). The active site of ficin consists of two amino acids:
Cys-25 and His-159, and up-to-date, only three fragments of ficin have been stud-
ied, (I) a fragment around the Cys catalytic site, (II) a His catalytic site, and (III) the
N-terminal fragment (Aider, 2021). Further, ficin has broad substrate specificity and
intense proteolytic activity toward the basic and neutral amino acids (Zhai
et al., 2021).
Ficin functions around a wide range of pH from 3 to 10 and the optimum activity
have been reported around neutral pH, generally around 6.5–8.5. It loses its activity
completely below pH 3, resulting in a partially folded structure (Devaraj et al.,
2008a, b; Milošević et al., 2019). The optimum temperature range for the ficins is
around 45–55 °C; however, its general activity could be observed from 30–90 °C
(Aider, 2021). Above 70 °C, ficins are prone to get denatured, and the apparent
denaturation temperature of ficin has been reported as 73 °C (Devaraj et al., 2008a,
b). According to Devaraj et al. (2008a, b), ficin loses its activity by 37% when incu-
bated for 10 minutes at 70 °C and loses 50% of its activity for 20 min at the same
temperature. The activity of the ficin enzyme has been reported to be enhanced in
the presence of cysteine and mercaptoethanol, whereas p-chloromercuribenzoate
and HgCl2 were identified as ficin enzymes inhibitors (Morellon-Sterling
et al., 2020).
The major problem associated with the ficins is the autolysis, which affects their
storage time and stability. According to Zare et al. (2016), the amount of peptide
production due to autolysis of ficin A, B, C, and the unretained fraction extracted
from F. carica cv. Sabz was increased when the storage time increased from 1 to
10 days. However, their autolysis rate differed significantly, and the ficin A and
unretained fraction showed much lower autolytic activities.

3.2 Other Proteolytic Components in F. carica

Besides Cys proteases, several other proteolytic components have also been identi-
fied from the F. carica latex. Hamed et al. (2020) discovered a serine protease
named FPIII, which has a specific activity of 2430 units/mg of protein. It showed
optimum activity around 8.5 pH and 60 °C. According to Hamed et al. (2020), the
relative activity of FPIII is highest on gelatin, following fibrin, albumin, hemoglo-
bin, casein, and collagen in the decreasing order. In addition to ficin, subtilase, per-
oxidase, chitinase, and mandelonitrile lyase are also present in F. carica latex
(Kitajima et al., 2018). A recent investigation on collagenolytic serine proteases and
chitinolytic enzymes present in fig latex has gained much attention for their exten-
sive proteolytic properties (Raskovic et al., 2016).
Further studies, however, are needed to configure the characteristics and proper-
ties of these fig latex-derived proteases. Raskovic et al. (2014) identified and puri-
fied another collagenolytic serine protease from the latex of F. carica variety Brown
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 753

Turkey using gel and covalent chromatography techniques. It is a highly stable

monomeric protease having a molecular mass of 41 kDa, showing activity around a
wide range of temperatures and pH. Further, this protease showed its specific activ-
ity toward gelatin and collagen. It possesses shallow non-specific proteolytic activ-
ity; hence, it is an excellent source to replace microbial collagenolytic enzymes in
industrial applications (Raskovic et al., 2014). Moreover a chitin-binding protease
with antimicrobial properties has also been identified and purified from the latex of
F. carica, which could be used in pharmaceutical and biotechnological industries
(Mavlonov et al., 2008).

4 Enzymatic Extractions and Sample Preparations

Pioneering studies on isolating different forms of ficin from fig latex were reported
in the early 1960s (Kramer & Whitaker, 1964). Ficin isolated from different culti-
vars of fig consists of closely related multiple isoforms (Milošević et al., 2019).
However, these ficin isoforms may have different conformational characteristics
due to the variation in the amino acid sequence in each isoform (Zare et al., 2013).
In addition, these ficin isoforms possess different isoelectric points, which facili-
tates their separation by chromatographic methods (Milošević et al., 2019). As
described earlier in this chapter, fig latex mainly contains an aqueous solution and a
gum portion (i.e., 70% water and 20% gum) (Zare et al., 2013). Therefore, the first
step of the isolation process is to remove the gum using a centrifugation process,
followed by dialysis to concentrate the supernatant (Kitajima et al., 2018). Ion-
exchange chromatography or cationic gel mediuma has been commonly used as a
separation technique to distinguish the different active fractions of purified ficin
(Azarkan et al., 2011; Zare et al., 2013). The main concern during the isolation
process of ficin is its stability concerning the structure and biological activity
(Baeyens-Volant et al., 2015). Zare et al. (2013) studied the homogeneity of the
peaks of the fractions eluted from the ion exchange chromatography using gel filtra-
tion using High-Performance Liquid Chromatography, electrophoresis using
sodium dodecyl sulfate-polyacrylamide gel electrophoresis, and recycling-
chromatography of the peaks using sulphopropyl-sepharose resins. Most of these
fractions have exhibited heterogenous peaks suggesting that ficin may have auto-
lyzed. Ficin A showed the lowest heterogenous peaks compared to other ficin iso-
forms, suggesting that ficin A is a suitable isoform for industrial applications.
Therefore, the use of autolysis inhibitors, i.e., iodoacetamide and potassium tetra-
thionate, could be a viable approach to maintain the stability of ficin during produc-
tion, isolation, purification, storage, and other industrial applications (Zare et al.
2013; Aider, 2021).
Early studies on fig lattices have used carboxymethyl (CM)-cellulose chroma-
tography to identify the proteolytic components of ficin based on its activity profiles
(Sgarbieri et al., 1964). However, such an approach may not provide detailed and
clear evidence for molecular mass information that may directly influence autolysis
754 H. Priyashantha et al.

(Azarkan et al., 2011). Disulfide bonds in proteases are resistant to autolysis and key
to protease stability. Therefore, it is recommended to use protective and reversible
measures to preserve free thiol groups present in ficin to avoid the adverse effects of
autolysis. The use of reversible inhibitors of the thiol group, including S-methyl
methanethiosulfonate or 2,20-dithiodipyridine, has been identified as an effective
method to produce simplified chromatograms. The thiol-pegylation strategy is
another practical approach to suppressing the autolysis and producing highly puri-
fied ficin forms (Azarkan et al., 2011). This includes rapid fractionation of ficin
isoforms followed by chemical modification using a monomethoxypolyethylene
glycol (mPEG) reagent (Azarkan et al., 2011). A recent study on ficin isoforms
investigated the potential of performing in situ chemical modification of the cata-
lytic cysteine to prevent the autolysis and other possible protein degradation of latex
proteases (Baeyens-Volant et al., 2015). This method includes collecting latex from
fig using methylmethanethiol sulfonate (MMTS) to prevent the possible denatur-
ation of ficin isoforms during processing (Baeyens-Volant et al., 2015). Mass spec-
troscopy is generally performed to characterize the de novo sequencing of purified
peptides to confirm the ficin isoforms (Azarkan et al., 2011). Moreover, Milošević
et al. (2020) reported that performing chromatography at an acidic pH (~5 pH)
could retard the autolysis process of ficin since cysteine proteases show lower activ-
ity at acidic pH values.
Furthermore, studying the comparative stability of cysteine proteases revealed
that the presence of multiple isoforms of ficin greatly influenced the stability of
papain. These findings suggested that ficin isoforms are more suitable for industrial
usages, i.e., therapeutics, diagnostics, bioreactors, biosensors, and fine chemical
applications, than other papain-like proteases (Milošević et al., 2019). Furthermore,
it was evident that ficin isoforms in the mixture could be more functionally stable
than isolated ficin isoforms (Milošević et al., 2020). In contrast to these findings,
separating ficin from other enzymes in the latex improved its effectiveness in pro-
teolytic activity. Finally, the complexity of ficin identification and characterization
may be associated with the genomic/sequential differences among varieties and cul-
tivars (Milošević et al., 2020). Thus, predominantly, ficin isoforms have been identi-
fied using approaches based on proteome and transcriptome analyses (Kitajima
et al., 2018; Aider, 2021).

5 Milk Clotting Properties of Ficins from F. carica

The capability of proteases to hydrolyze the κ-casein is referred to as the milk clot-
ting activity (Jacob et al., 2011). It is an essential parameter in determining the suit-
ability of protease in cheese-making. The milk clotting index is the ratio between
milk clotting activity and proteolytic activity of proteases. Higher the milk clotting
index, higher the suitability as a rennet substitute since it possesses high milk clot-
ting activity with low general proteolytic activity. High proteolytic activity is the
major drawback of plant proteases since it leads to excessive hydrolysis of milk
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 755

caseins resulting in bitter flavors, textural defects, and yield loss of cheese (Roseiro
et al., 2003; Jacob et al., 2011; Anusha et al., 2014; Shah et al., 2014). Although
numerous plant proteases have been proposed as rennet substitutes, they do not pos-
sess a high enough milk clotting index to use at the industrial level (Anusha
et al., 2014).
Among several proteolytic components in the fig latex, only ficins have been
reported to have milk clotting activity (Raskovic et al., 2016) and visualized in
Fig. 33.3, using crude extract of fig. Therefore, despite their high proteolytic activ-
ity, numerous attempts have been made to produce various cheese types from differ-
ent types of milk, using ficins as milk coagulants. Bornaz et al. (2010) studied the
proteolytic activity of ficin, cardosin, and chymosin on fresh cheese production
using ewe’s milk. According to their findings, ficin resulted in a great amount of
curd loss in the whey, which is a negative perspective regarding cheese-making, and
the total solids, total nitrogen, and the casein amount in the whey also appeared to
be highest when ficin is used. However, despite the significant loss of caseins, ficin
yielded the highest cheese yield due to its high water retention ability and low syn-
eresis capacity compared to cheese obtained from cardosin and chymosin.
Nevertheless, no significant differences in sensory properties were observed among
the three types of curds (Bornaz et al., 2010). Daffri et al. (2019) compared the milk
clotting activity of ficin from two F. carica varieties: Marseillaise and Dauphine,
and found that both types possess similar milk clotting activity (1100 UAC) even
though the Marseillaise variety had higher proteolytic activity than Dauphine.
Raskovic et al. (2016) reported the changes in the protease profile of fig latex
over the fruit ripening process. According to their findings, the ficins concentration

Fig. 33.3 Milk clotting ability of F. carica. Raw milk (a) is coagulated (within 30 min) into a
semi-solid coagulum (b) using crude extract of F. carica whole fruit. (Photo: Hasitha Priyashantha)
756 H. Priyashantha et al.

in the latex altered over time. At the beginning of fruit ripening, fig latex contains
high concentrations of ficin forms, which possess high caseinolytic activity, leading
to a low milk clotting index. The authors predicted that, with the fruit ripening, ficin
isoforms with broad specificity for caseins are gradually replaced with isoforms
having high specificity towards κ-casein. Therefore, the authors suggested that the
ficins derived from ripened fruits or harvested in summer are the best source for
milk coagulation as they possess high milk clotting activity and milk clotting index
(Raskovic et al., 2016). An early study by Kamer & Whitaker (1964) also reported
the milk clotting properties of ten isoforms of ficins derived from F. carica var.
Kadota found that different isoforms have different specificity toward caseins.
According to their findings, all ten isoforms showed caseinolytic activity, and
among them, the isoforms termed B and D showed more or less similar and the
highest caseinolytic activity. All the isoforms except isoform A showed milk clot-
ting activity; however, only the isoforms C, D, E, and G resulted in considerably
higher milk clotting activity than other isoforms. Moreover, the authors reported
that the isoforms C, D, E, F, and G are more heat and pH-stable than isoforms.
Among them, isoform G appeared to be the best form for milk coagulation, as it
possessed comparatively lower caseinolytic activity and higher milk clotting activ-
ity, resulting in the highest milk clotting index of all other isoforms (Kamer &
Whitaker, 1964).
Akar and Fadiloglu (1999) studied the properties of a “Teleme” cheese and found
that the Teleme cheese made with purified ficin had better chemical and sensory
properties than that made with crude fig latex. This might be due to removing non-
casein-specific proteases with the purification process. According to their study, the
purified ficin using ion-exchanged chromatography gave about two-fold higher
milk clotting index than the crude latex. Moreover, further purification of ficin using
gel filtration resulted in about four times higher milk clotting index than the crude
latex. However, purified ficins took much longer to coagulate milk than crude latex.
A study of using ultra-filtered bovine skim milk in cheese production revealed that
ficin performed better in more concentrated milk (4 times ultra-filtrated) than regu-
lar milk, giving less free amino acids and less clotting time and firmer curd. However,
ficin resulted in lower cheese yield than other proteases (cardosin and chymosin),
making it unsuitable for bovine milk, especially when using regular milk (Low
et al., 2006). Siar et al. (2020) reported that ficin immobilized using glyoxyl-agarose
led to dense coagulum and gave a higher yield than the free-ficin when coagulating
skim milk. For that, milk coagulation should separate into two steps, (I) enzymatic
hydrolysis of casein at 4 °C to prevent hydrolysate aggregation, which causes sepa-
ration of the aggregate and biocatalysts, and (II) force hydrolysate aggregation by
heating up to 40 °C. According to the authors, this method increases clotting yield
by up to 27% (Siar et al., 2020). Ficin activity on camel milk has been studied by
Fguiri et al. (2021), and according to their findings, ficin gave more acidic cheese
rich in fat, dry matter, ash, and proteins than the chymosin. Moreover, ficin resulted
in higher microbiological quality in cheese made with camel and cow milk than
chymosin (Fguiri et al., 2021).
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 757

Overall, ficin appeared to be more suitable in coagulating ovine and caprine milk
than bovine milk, as ficin resulted in poor milk clotting properties in bovine milk.
However, ultrafiltration of milk could enhance the activity of ficins; as the concen-
tration of milk increases, the mean distance between casein micelles decreases,
resulting in increment of aggregation velocity and reduced clotting time, thus lesser
milk protein hydrolysis. Moreover, purification of ficin often leads to better coagu-
lation properties. Current knowledge of the activity of ficins against coagulation of
different milk types facilitates a clear understanding of the potential and limitations
of using ficins as a commercial calf rennet replacer. Nevertheless, further studies
will enhance its precise usage in milk coagulation as a cheap yet, efficient milk
coagulant.

6 Other Beneficial Roles of Using F. carica

in Milk Coagulation

Ficin has great potential for use in commercial applications and is considered an
emerging plant-based enzyme due to its strong catalytic activity. In particular, ficin
has been successfully used in the dairy industry as a coagulant for the cheese manu-
facturing process as an alternative to animal-originated coagulants (Aider, 2021). In
addition to the coagulant activity, ficin has exhibited unique proteolytic activity in
the presence of milk proteins (Baeyens-Volant et al., 2015). This characteristic fea-
ture of ficin has the potential to produce milk protein hydrolysates as a functional
food ingredient (Milošević et al., 2020). The milk protein hydrolysis process results
in an array of bioactive peptides that can be used as functional food ingredients for
promoting health and reducing the risk of certain diseases (Rasika et al., 2015;
Abdel-Hamid et al., 2017). Bioactive peptides are generally encrypted within the
precursor protein structure as inactive peptide fragments that can be released in vivo
or in vitro enzyme hydrolysis (Rasika et al., 2015; Abdel-Hamid et al., 2017). Ficin
exhibits consistent and rapid proteolytic hydrolysis maps of proteins compared to
other plant-derived enzymes (Siar et al., 2020). This specific characteristic of ficin
has been extensively studied in milk casein protein to determine the bioactive poten-
tials of resulting peptides. Antioxidant activity was one of the most investigated
bioactive properties in food-derived peptides. Antioxidant peptides can interact with
the free radicals and reactive oxidative species to prevent or retard the oxidation
process in biological materials and food systems (Shahidi & Zhong, 2008). Potent
antioxidative peptide sequences have been identified from bovine casein hydroly-
sates produced using F. carica L. derived proteinase (Di Pierro et al., 2014). The
radical scavenging activity of produced peptides was evaluated using an oxygen
radical absorbance capacity (ORAC) assay that quantified the percentage and degree
of inhibiting free radicals. F. carica L. latex proteinase has the potential to hydro-
lyze casein substrate and produce low molecular weight peptides that may be
responsible for its radical scavenging ability (Di Pierro et al., 2014).
758 H. Priyashantha et al.

In addition to antioxidant activity, antimicrobial activities of milk protein hydro-

lysates produced from ficin have been reported in recent literature. Goat milk casein
was hydrolyzed using ficin and trypsin and resultant peptides were analyzed for
their inhibitory activity against representative Gram-positive and Gram-negative
bacteria (Esmaeilpour et al., 2016). Among the enzyme treatments, extensive prote-
olysis of goat milk substrate was observed with ficin treatment. Moreover, the high-
est antimicrobial activity against E.coli was reported in ficin-assisted hydrolyzed
samples. The study’s findings further demonstrated that peptides composed of
hydrophobic and charged (polar) residues exert excellent antimicrobial activities.
This indicates that ficin could be a promising plant-derived enzyme to produce
potent antimicrobial peptides (Esmaeilpour et al., 2016). Furthermore, Raskovic
et al. (2016) investigated the antifungal activity of fig latex and its proteolytic and
milk clotting properties. They reported that different isoforms of ficin with various
casein specificities were present in the fig latex during different fruit ripening
phases, and the highest antifungal activity of ficin was exhibited in the early phases
of flowering (Raskovic et al., 2016). The authors also indicated that F. carica latex
has the potential to be utilized as a natural food preservative and a functional food
ingredient due to its unique compositional characteristics, including antioxidant
compounds, vitamins, minerals, fibre, and proteins (Raskovic et al., 2016).
Shabani et al. (2018) studied the rheological and microstructural characteristics
of ewes’ milk curd obtained by ficin and Polyporus badius enzymes. They reported
that increasing the concentration of ficin has a pronounced effect on gel elasticity
and other beneficial microstructural properties (Shabani et al., 2018). Furthermore,
the use of latex peptidases in reducing the allergenicity of milk proteins has also
been investigated to develop novel hypoallergenic formulas for children with a milk
allergy (Oliveira et al., 2019). In this regard, ficin could be a promising candidate for
producing extensively hydrolyzed milk proteins with reduced antigenicity and aller-
genicity (Abd El-Salam & El-Shibiny, 2017). This characteristic feature of ficin is
closely linked with its proteolytic activity, which can adversely affect IgG or IgE
antibody-mediated responses (Aider, 2021). However, further clinical studies are
recommended to illustrate the mechanism of action related to the immunomodula-
tory activity of milk protein hydrolysates derived from ficin. Enzymatic hydrolysis
of milk proteins is also considered an effective approach to enhancing bioactive
peptides’ bioavailability and beneficial amino acids. The produced milk hydroly-
sates have gained much attention as a key ingredient in food supplements, infant
formula, sports nutrition, therapeutic diets, and geriatric nutrition (Abd El-Salam &
El-Shibiny, 2017; Aider, 2021). However, up-to-date, limited studies have been con-
ducted to determine the sensorial properties of milk protein hydrolysates produced
from plant-derived latex proteases. Thus, extensive investigations of the distinct
proteolytic activity of ficin on milk proteins will pave the pathways to identify its
diverse applications in functional food, nutraceutical, and pharmaceutical industries.
33 Use of Proteolytic Activity of Ficus carica in Milk Coagulation 759

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Chapter 34
The Potential of Fig (Ficus carica) for New
Products

Sara Khoshnoudi-Nia , Akram Sharifi , and Elham Taghavi

1 Introduction

The fig (Ficus carica L.) is a tree or shrub with a pear-shaped and fleshy fruit from
the Moraceae family. Fig is native to southwestern Asia and the eastern part of the
Mediterranean due to suitable climate conditions (Hossain et al., 2010; Mawa et al.,
2013). Fig is a very delicious fruit and rich in fiber, proteins, amino acids, carbohy-
drates, sugars, minerals (copper, manganese, magnesium, potassium, and calcium),
organic acids, vitamins, and bioactive compounds (e.g., antioxidant polyphenols,
flavonoids, anthocyanin pigment) which are the reasons could explain the growing
demand for this fruit in the world (Petkova et al., 2019; Shamin-Shazwan et al.,
2019). Furthermore, due to the high concentration of calcium (133 mg/100 g; a
higher than 10% of the RDI or recommended dietary intake) of fig fruits, figs could
be considered “rich in calcium” (Barolo et al., 2014).
Moreover, the phytochemicals have been found in figs as follows: arabinose,
β-amyrins, β-carotenes, glycosides, β-sitosterols, xanthotoxol, alkaloids, flavonoids
(e.g., rutin, catechin, and epicatechin), coumarins, phenolic acid (chlorogenic acid,

S. Khoshnoudi-Nia (*)
Seafood Processing Research Group, School of Agriculture, Shiraz University, Shiraz, Iran
e-mail: [email protected]
A. Sharifi
Department of Food Science and Technology, Faculty of Industrial and Mechanical
Engineering, Qazvin Branch, Islamic Azad University, Qazvin, Iran
e-mail: [email protected]
E. Taghavi
Faculty of Fisheries and Food Science, Universiti Malaysia Terengganu,
Terengganu, Malaysia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 765
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_34
766 S. Khoshnoudi-Nia et al.

hydroxycinnamic acids (3-O- and 5-O-caffeoylquinic acids, ferulic acid, gallic acid
and syringic acid), flavonoid glycosides (quercetin-3-O-glucoside and quercetin-3-
O-rutinoside), furanocoumarins (psoralen and bergapten), saponins, and terpenes
(Çalişkan & Polat, 2011; Petkova et al., 2019; Pourghayoumi et al., 2016; Tanwar
et al., 2014; Veberic & Mikulic-Petkovsek, 2016). Because of these natural bio-
compounds, several health benefits have been reported for fig fruit, and it has an
important place in the Mediterranean diet. For example, fresh and dried figs are
recommended for enhancing eyesight, liver, and spleen diseases. In addition, it
could be used as a good laxative, expectorant, and diuretic agent (Badgujar et al.,
2014; Shahrajabian et al., 2021).
Fig is consumed either in fresh or dried fruit. Fresh Fig has a short post-harvest
life (around one week) and is highly sensitive to microbial activity due to high mois-
ture content. A combination of modified atmosphere packaging and cold storage
conditions may extend this time up to 2–4 weeks (del Carmen Villalobos et al.,
2016; Villalobos et al., 2016). However, the drying procedure can reduce the micro-
bial load of fruits by decreasing the water activity and consequently minimizes
physical and biochemical changes during storage. Therefore, figs are commercially
available in dried form. The annual production of dried figs is about 1.332813 mil-
lion tons in Turkey (310,000 t), Egypt (225,295 t), Morocco (153,472 t), Iran
(130,328 t), Algeria (114,092 t), and Spain (51,600) are the main producer of this
fruit (FAOSTAT, 2020). In most fig-producing countries, fig fruit is sun-dried to
save energy and cost. Moreover, the sun-drying method is an environment-friendly
process. The suitable fresh fig fruit for drying is at full maturity and after wrinkling
and dehydrating (over-ripe stage) on the tree (Aksoy, 2015).
Fresh or dried fig fruit can be processed to produce varied food products such as
jam, jelly, fig powder, marmalade, wine, liquor, juice, etc. Moreover, several by-
products (such as low-quality figs and wastes of juice) are generated during fig
processing, which is a valuable source of bioactive compounds like phenolic com-
pounds, colorants, sterols, etc. Therefore, the current chapter aims to discuss several
fig products and by-products, evaluate the influence of the food processing methods
and storage conditions on antioxidants, and review the nutritional composition of
fig products and by-products. Furthermore, recent findings on the potential of figs to
develop new functionality and nutraceutical products will be highlighted.

2 Fig Products

2.1 Dried Fig

One of the main forms of consumption of figs is dried form. Fig fruits are usually
dried by a natural sun drying method. However, artificial drying systems can also be
used. The energy cost is the most significant factor in selecting an artificial tech-
nique. In addition, the fig quality and maturity, drying time and temperature regime,
34 The Potential of Fig (Ficus carica) for New Products 767

airflow rate, and relative humidity affect the final dried fig quality (Freiman et al.,
2015). Fig fruits with thin and fleshy skin containing high sugar and low acidity
level are the most suitable fresh fig fruit for drying, and these dried figs are softer
and sweeter. Fresh fig fruit can be peeled and sun-dried to produce dried fruits with
a softer texture and lighter color. Also, fresh fruit may be sliced or cubed before
drying (Aksoy, 2015). Cut or cubed dried figs mixed with nuts and cereals for pro-
ducing snacks, muesli, and granola. The cubed figs may be coated with chocolate
(Isa et al., 2020).
The bioactive compound content of fresh or processed fig fruit strongly depends
on the cultivar type (De Pilli et al., 2019). Khradhraoui et al. (2019) investigated the
phytochemical content, antioxidant potential, and fatty acid composition of nine
sun-dried fig cultivars originating from South-Eastern and Middle Eastern Tunisia.
Dark fruits exhibited a higher total polyphenol content (Saoudi Douiret cultivar:
201 mg GAE 100 g−1 dm) than green ones (Bayoudhi Douiret cultivar: 73.74 mg
GAE 100 g−1 dm). Fatty acid methyl esters contained (C16: 0), (C18: 1), ((C18: 2)
9, 12), ((C18: 3) 9, 12, 15) and (C20: 0). Strong correlations were found between the
amounts of various phenolic compounds and fatty acids with antioxidant capacity
(Khadhraoui et al., 2019).
The method of the drying process may affect the nutritional and health-related
properties. In this regard, Slatnar et al. (2011) showed that the highest sugar/organic
acid ratio was obtained for the sun-dried sample. Furthermore, the higher content of
all individual phenolic compounds (chlorogenic acid, catechin, epicatechin,
kaempferol-3-O-glucoside, luteolin-8-C-glucoside, rutin, quercetin-3-O-glucoside,
and cyanidin-3-O-rutinoside) was determined after the oven-drying process.
Similarly, higher total phenolic content and antioxidant activity were detected after
the oven drying process. The results indicated that dried figs are a good source of
phenolic compounds and organic acids. However, these authors expressed the nutri-
tional properties based on moisture weight, not dried ones (Slatnar et al., 2011).
Moreover, Vallejo et al. showed that after fig drying, the losses of polyphenols did
not exceed 15% (Vallejo et al., 2012).
Not only is the quality important, but safety is an essential factor in fig dried.
Many studies reported the risk of the presence of mycotoxins in dry fruits, espe-
cially in dry figs (Sulyok et al., 2020; Trucksess & Scott, 2008; Yujiao Wang et al.,
2018). Aflatoxins are considered the most toxigenic metabolites of the mycotoxin
classes produced mainly by Aspergillus flavus and Aspergillus parasiticus (Ait
Mimoune et al., 2018). Treating the fresh fig with Sulphur before drying caused a
lighter color and prevented pest damage. The maximum residue limit for SO2 in
dried figs is 1000 ppm (Codex Alimentarius, 1989). Moreover, the labeling of food
containing sulfites (>10 ppm) is obliged by US Food and Drug Administration
(FDA). Methyl bromide was the main fumigant for controlling pests before 2015.
However, it has been banned in developed and developing countries since 2005 and
2015. To obtain sun-dried figs, figs are exposed to sunlight to complete the dehydra-
tion process, increasing fungal contamination and aflatoxins development (Ait
Mimoune et al., 2018). A controlled drying process helps to reach a safety level to
enhance the protection of public health. Petrić et al. (2018) treated the figs with
768 S. Khoshnoudi-Nia et al.

various solutions as followed: 0.5% citric acid; 0.5% ascorbic acid; 0.3% L-cysteine;
0.2% chestnut extract; 0.15% Echinacea extract before drying in a cabinet dryer.
They found nine metabolites (i.e., aflatoxin B (AFB1), ochratoxin A, ochratoxin
alpha, kojic acid, emodin, altenuene, alternariol methyl ether, brevianamide F, and
tryptophol in fig samples. However, all pretreatment solutions reduced AFB1, being
a major fig contaminant. The most efficient pretreatment was L-cysteine (15%
reduction), while ascorbic acid increased mycotoxin (158% increase). However, all
pretreatment solutions reduced AFB1 as a major fig (Petrić et al., 2018). Ozone is
the green and effective alternative for Methyl bromide (Akbas & Ozdemir, 2008;
Sadeghi et al., 2017).

2.2 Fig Paste and Puree

Dried fig fruit that is not fit for direct human consumption was classified as Class 2
or substandard or surplus. These figs are processed to develop new products,
decrease waste and make value-added. These figs may be ground into a paste. In fig
paste production, the seeds can be crushed or not, depending on the demand. The fig
paste or puree can be used to fill cookies and cakes and as a raw matter in making
ice cream, jam, marmalades, or juice. Fig bars and fig balls also may be prepared
with fig paste. Small fig balls can also be coated with chocolate or mixed with vari-
ous nuts and then coated with roasted sesame seeds, chia seed, coconut, or cocoa
powder (Aksoy, 2015).

2.3 Fig Jam

Jam production is one of the most popular ways to enhance shelf-life and preserve
fig fruit. The steps of jam processing were as follows: the fresh figs were sorted,
washed, and cut (the fruit can also be used without cutting). For the inactivation of
the enzymes, the fruits were heated at 80 °C. Then, sugar dissolved in water was
added and boiled to reach a final soluble solid concentration (600–700 g/kg). Then,
pectin was added, and the mixture was heated for 10 min to complete the hydration
of the pectin. Finally, hot-packing (85 °C) and sterilization were conducted.
Although jams are popular fruit products, they contain high levels of sugar. Instead,
preserves are produced without additional sugar being better alternatives for con-
ventional jams (Levaj et al., 2010).
Jam formulation affects the physicochemical, nutritional, and sensory properties
of fig jams. Kumari et al. (2018) evaluate the effect of various concentrations of
pectin (0.3–0.7%) and citric acid (0.3–0.7%) on physicochemical and sensory prop-
erties of wild Himalayan fig jam. The fig jams contained 0.7% pectin, and 0.3%
citric acid was the best formulation. This jam showed higher titratable acidity
(12.0%: based on citric acid), ascorbic acid content (1.24 mg/100 g), total sugar
34 The Potential of Fig (Ficus carica) for New Products 769

(66.4%; reducing sugar: 58.8%), total phenol (39.8 mg/100 g), anthocyanin
(17.0 mg/100 g) and sensory scores. The FTIR curve of fig jam and its pulp showed
no alteration in physicochemical parameters during jam processing on fig pulp
(Kumari et al., 2018).
The ripeness of fig fruit can also affect the properties of the jam. Levaj et al.,
2010 evaluated the gel strength and sensory properties of jams and preserves made
from figs with different ripeness. During ripening, the total solids, sugars, and pH
values were increased in jams and preserves. Investigation of the amount of three
pectin fractions (soluble in water, soluble in oxalate, and alkali) showed that the
most abundant pectin fraction of figs was soluble in oxalate one. It was not affected
by the stage of ripeness. The amount of soluble in water and soluble in alkali frac-
tions (pectin fractions responsible for the strength of gels) decreased with increas-
ing fruit ripeness. Gels made soluble in oxalate pectin were significantly softer than
other gels. While the ripeness stage had not affected the strength of jams, with
increased fruit ripeness, the preserve’s strength decreased. The sensory properties
(on taste, firmness, and color) of jams or preserves were not affected by the stage of
ripeness. However, the sensory scores of jam were higher than preserves (Levaj
et al., 2010).
Furthermore, the literature review showed that the bioactivity of fig fruit is
affected by processing conditions. Tanwar et al. (2014) also evaluated the effect of
jam and nectar processing on fig fruit products’ physicochemical and nutritional
properties. The results revealed that conversion of fig fruit pulp into jam and nectar
caused a significant increase in total soluble solids and titratable acidity, carbohy-
drate, and energy value but a decrease in pH, iron, calcium and phosphorus, mois-
ture, ash, crude fiber, crude protein, crude fat, vitamin C, ß carotene, total phenolics,
total flavonoids, total anthocyanins and tannins (Tanwar et al., 2014). Moreover,
these authors (2015) assessed the effect of similar processes on the antioxidant
activity of fig fruit. Fig pulp showed higher antioxidant activity than fig jam and fig
nectar. Total phenolics decreased by 10 and 55%, flavonoids by 98 and 45%, antho-
cyanins by 79 and 33%, and tannins by 83 and 77% in fig jam and fig nectar as
compared to fig pulp, respectively (Tanwar et al., 2015).
The effect of cold storage and jam processing on physicochemical properties and
phytochemical compounds of yellow-colored figs (Ficus carica L.) has been evalu-
ated by Petkova et al. (2019). The highest and lowest bioactive compounds content
and antioxidant activity were found in fresh and frozen fruits, respectively (after
4 months of storage at −18 °C) The jam processing was a better technique to pre-
serve bioactive compounds as compared freezing method (especially for carot-
enoids and phenolic compounds). After the jam and freezing process, the losses of
total phenolic content were around 10% and 45%, respectively (Petkova et al.,
2019). Rababah et al. (2011) found that total phenolics and anthocyanins of fig jam
decreased by 68.6% and 60.2% after 5 months of storage. Despite reducing these
compounds in jam processing, it is a good method to enhance the shelf-life of
fig fruit.
Abd-El-Hak et al. (2016) blended the sycamore and fig fruit to formulate a nutri-
tional and value-added jam. Sycamore and fig fruit were blended in the ratios of
770 S. Khoshnoudi-Nia et al.

100:0, 0:100, 20:80, 80:20, and 50:50, respectively. Various jam samples’ chemical
and sensory properties were evaluated during 12 months of storage at room tem-
perature. The results showed that sycamore was rich in ash and crude fiber. The
sensory score of jam samples was improved by increasing the fig content of jam up
to 50%. During storage, the acidity of samples decreased, and their physiochemical
properties improved. Therefore, the blend of sycamore and fig in jam production
was recommended to increase nutritional value.

2.4 Fig Juice

As a healthy drink, Fig juice is an excellent source of vitamin C and natural poly-
phenols (e.g., flavonoids and proanthocyanidin), helping boost the immune system
and balance blood pressure (Joseph & Raj, 2011). preparation of fig juice is done
directly by applying pressure to fresh fig or boiling the dried fig in water and extract-
ing the procedure. Soft and juicy fresh figs are more suitable for obtaining fig juice
directly. Furthermore, fig juice and concentrate can be consumed directly or used in
the pastry industry as glazing and bio-sweetener (Soni et al., 2014).
Although fig juice contains a high level of phenolic compounds, most of these
compounds are difficult to be absorbed by the human body. To improve the absorp-
tion, hydrolysis by enzymes or microbes in the digestive tract is a suitable solution
(Filannino et al., 2016). Fermentation can enhance the nutritional value and antioxi-
dant activity of food products. Various biochemical changes (especially conversion
of glycoside into the aglycone and increasing the flavonoid release) occurred during
the fermentation causing the nutrition component and anti-nutrition ratio bioactivity
and digestibility changes (Zhang et al., 2012). On the other hand, beverages are an
appropriate delivery system for probiotics. There are several health benefits related
to the probiotics, such as antimicrobial, antimutagenic, anticarcinogenic, antidiar-
rheal, and antihypertensive properties, enhancing lactose digestion, reducing cho-
lesterol and mineral malabsorption Therefore, probiotic fig juice can be considered
a healthy and nutraceutical beverage for vegetarians and consumers with lactose-
allergy (Khezri et al., 2016). Khezri et al. (2016) compared the survivability of three
species of lactic acid bacteria (Lactobacillus casei, L. plantarum, and L. delbrueckii)
in fig juice. Heat-treated fig juices were treated by three species (6 log CFU mL−1)
separately and incubated at 30 °C for 72 h. Among various species, L. delbrueckii
showed the best survivability in high acidity and reached nearly 9 log CFU mL−1
after 48 h of fermentation. After 4 weeks of cold storage (⁓4 °C), the viable cell
counts of L. delbrueckii and L. plantarum were 6 and 5 log CFU mL−1, respectively.
However, L. casei was just survived for 2 weeks. Therefore L. delbrueckii was intro-
duced the most suitable strain among other species at the consumption time.
Therefore, probiotic fig juice can serve as a healthy beverage for vegetarians and
consumers with lactose allergies (Khezri et al., 2016).
Wijayanti et al. (2017) used Lactobacillus acidophilus, Lactobacillus bulgari-
cus, Lactobacillus casei, and Lactobacillus plantarum as a starter (at 37 °C for 24 h)
34 The Potential of Fig (Ficus carica) for New Products 771

for producing fermented fig juice. The fermented fig juice demonstrated higher total
phenolic content. L. bulgaricus showed the highest increase in total phenolic con-
tent (0.45%) and antioxidant activity (IC50 = 76.55 ppm) as compared to the control
ones (Total phenolic compound = 0.09%; IC50 = 76.7 ppm) (Wijayanti et al., 2017).
To enhance the viability of probiotics in an acidic medium, Khezri et al. (2018)
formulated synbiotic fig juice. They used L. delbrueckii as a probiotic culture and
inulin as a prebiotic ingredient and evaluated the effect of various formulations on
microbial survivability, physicochemical, total phenolic content, antioxidant capac-
ity, and sensory parameters of fig juices. After 48 h of fermentation, the cell contents
of lactic acid bacteria reached the maximum level in both probiotic (8.41 Log CFU
mL−1) and symbiotic (8.83 Log CFU mL−1) samples. The total phenolic content and
antioxidant capacity of fermented fig juices were significantly higher than the con-
trol samples. The viability of Lactobacillus delbrueckii significantly decreased dur-
ing storage time. However, the rate of decline was slower in the synbiotic juice (7.49
Log CFU mL−1 after 4 weeks of cold storage) than in the probiotic one (6.59 Log
CFU mL−1). The highest overall sensory score was obtained for control juice (7.8),
followed by synbiotic juice (6.1) and probiotic juice (5.7). Synbiotic fig juice was
better in terms of probiotic and antioxidant properties. At the end of the storage
period (4 weeks), the viable count of Lactobacillus delbrueckii in synbiotic samples
remained about 1 Log CFU mL−1 higher than probiotic ones being above the mini-
mum requirement (6 Log CFU mL−1) for probiotic food products (Khezri et al.,
2018). Overall, fig juice could be a suitable carrier for probiotics, and prebiotics and
fermentation could enhance the bioavailability of polyphenols.

2.5 Fig Jelly

To obtain the fig jelly, fig pulp, sugar, around 0.5% citric acid, and 1% pectin were
mixed. The fruit mixture was heated until boiling and reaching ⁓65 Brix. The fin-
ished product was pasteurized in jars with lids (Bof et al., 2012; Curi et al., 2019).
Fig variety can be affected the nutritional and physicochemical characteristics of fig
jelly. Therefore, Curi et al. (2019) assessed the effect of fig cultivars on the proper-
ties of fig jelly. The results showed that the various fig cultivars obtained jellies with
different physical, chemical, and rheological properties. However, the fig cultivar
had no significant effect on the acceptability of fig jelly (Curi et al., 2019). The
effects of freezing and jelly processing on the antioxidant capacity of various fruit
(e.g., grape, apple, strawberry, pear, guava, and fig (Ficus carica L.) for 90 days was
also investigated by Bof et al. (2012). Freezing (−15 °C) did not significantly differ
in the antioxidant capacity of grape and fig pulp. However, fig pulp showed the low-
est antioxidant activity. This lower antioxidant activity may refer to the variety and
harvest dates. However, the antioxidative compounds of strawberry and fig jelly
were heat-resistant and antioxidant activity remained stable in fig jelly (Bof
et al., 2012).
772 S. Khoshnoudi-Nia et al.

2.6 Wine and Vinegar

Wine is an alcoholic beverage (alcohol content: 5 to 13%) made from fermented

fruits. The high sugar content of fig fruit makes it a good candidate for producing
wine and vinegar. Yeast converts the juice sugar to ethanol and carbon dioxide
(Sheng et al., 2019). Fruit wine contains most of the fruit nutrients, alcohol, vitamin,
and other compounds produced by the yeast activation that moderate consumption
of it may protect people against certain cancers and improve mental and heart health
(Artero et al., 2015).
The properties of fruit (fresh or dried fruit, pH values, sugar, nitrogen contents,
etc.), type of yeast, and fermentation conditions can affect the wine quality (Chilaka
et al., 2010). The juice was mixed with 0.05% Diammonium hydrogen phosphate
(DAHP) and pasteurized at 85 °C for 30 min. Pasteurized juice was inoculated with
Saccharomyces cerevisiae (at 24 °C temperature for 20 days). The wine properties
are as follows: Brix: 8.5°; TSS: 0.4 to 4.2%; total sugar: 0.2 to 4.0%; alcohol: 9.9 to
11.8% alcohol. During the fermentation period, total soluble solids, titratable acid-
ity, reduced sugar, total sugar decreased, and the alcohol percentage of wine
increased. They introduced the wine prepared at 10% inoculums and 24 °C tem-
perature for 20 fermentation days as the best quality sample (PotdarVrushali
et al., 2020).
The fig processing conditions can be affected the quality of the wine. For exam-
ple, Lu et al. (2021) reported that dried fig wines had lower anthocyanin contents
and antioxidant activity than wines prepared with fresh samples due to the thermal
degradation of anthocyanins during the drying process. However, the drying process
had no significant effect on the total polyphenol and flavonoid contents (Lu
et al., 2021).
For wine production processes, one of the most important steps is the choice of
yeast. Since Saccharomyces showed a relatively high fermentation rate and yield
(alcohol production), it is still the most frequent yeast in wine production (Lu et al.,
2021). However, it was shown that non-Saccharomyces might improve the flavor
complexity of wines by producing various desirable volatile compounds (Binati
et al., 2020; Hranilovic et al., 2018). Liu et al. (2021) evaluated the quality of fig
wines co-fermented by three non-Saccharomyces (Wickeramomyces anomala;
Hanseniaspora uvarum, and Pichia fermentans). The results showed that fig wines
fermented by non-Saccharomyces exhibited higher contents of organic acids and
mono-phenols. Furthermore, co-fermentation by non-Saccharomyces could obtain
up to 38 aroma compounds, while fermentation by Saccharomyces obtained around
30 substances. Therefore, the aroma of the different samples showed significant dif-
ferences. The result revealed that pleasant fig wine could be obtained by co-
fermentation of non-Saccharomyces without a Saccharomyces starter culture (Liu
et al., 2021).
34 The Potential of Fig (Ficus carica) for New Products 773

3 Value-Added Fig Products

By increasing the world’s population and the global food demand, the food industry
and agriculture sector have increased their production. As a result of the increased
production and consumption, a considerable amount of agro-food waste and by-
products are produced, which are generally left unused and unattended (Sharma
et al., 2020). Therefore, it is crucial to establish ways to recycle these valuable
wastes. They can be used as sustainable and renewable resource materials for syn-
thesizing cost-effective, value-added products for various applications. In addition,
the products are in line with global health, food security, and sustainable agriculture
strategies (Dey et al., 2021).
Fig fruit by-products (peel, seeds, and no-optimal fruits) are generated during fig
processing to produce fig jam, fig juice, wine, etc. These by-products contain vari-
ous bioactive compounds and functional ingredients. Recycling these by-products
to produce individual compounds and/or extracts and additives for the food industry
can improve food products’ functional, nutritional, and techno-functional character-
istics (Teruel-Andreu et al., 2021). In this sub-section, some of the value-added
products obtained from the peel, seed, and fig pulp were reviewed.

3.1 Extraction of Phenolic Compounds

Phenolic compounds are important constituents of fruits that contribute to the taste,
color, and human health effects fruits (Khoshnoudi-Nia et al., 2020; Veberic et al.,
2008). Phenolic compounds and carotenoids are considered the most important
natural antioxidants (Tsao & Deng, 2004). Fig by-products are a rich source of
phenolic compounds. Phenolic acids (e.g., chlorogenic acid, syringic acid, caffeic
acid, ferulic acid, coumaric acid, quinol, and gallic acid) and flavonoids (e.g. (+)-cat-
echin, (−)epicatechin, anthocyanin, kaempferol, quercitin, and myricetin) are the
two major classes of fig fruits phenols (Bucic-Kojic et al., 2011; Dueñas et al.,
2008). In this regard, based on the conventional and modern isolation and character-
ization methods, 126 chemical constituents in the eight major groups (hydroxyben-
zoic acids, hydroxycinnamic acids, flavonoids, coumarins, furanocoumarins,
volatile constituents, triterpenoids, and miscellaneous) were found for Ficus carica
(Badgujar et al., 2014). Several phenolic compounds were found in fig fruit. The
type and level of phenolic compounds can be varied depending on the variety, part
of the fruit, time of harvest, maturity level, and extraction conditions (Vallejo et al.,
2012). Vallejo et al. (2012) collected 18 fig cultivars commonly grown in south-
eastern Spain in spring and summer and evaluated their polyphenolic profile.
Depending on the season crop, the total phenolic compounds ranged between 19.1
to 140 mg/100 g. Fruit harvested in spring generally showed higher phenolic values
than samples obtained in summer. Flavonoids are considered “environmental com-
pounds”. Because phenolic compounds were directly produced in response to
774 S. Khoshnoudi-Nia et al.

environmental conditions (such as ultraviolet light and CO2 levels) (Caldwell et al.,
2005). Moreover, Vallejo et al. (2012) reported that a high concentration of phenolic
compounds is present either in the skin (mainly anthocyanins, especially cyanidin-
3- rutinoside) or pulp (mainly proanthocyanidins, especially (epi)catechin) of fig
(Vallejo et al., 2012). Analyses of the potential health-promoting compositions of
10 Spain cultivars of fig fruits showed that the phytochemical profiling of fig varied
based on cultivar and the varietal type. In fig fruit, eleven polyphenolic compounds
(in flavan-3-oils, phenolic acids, flavonols, flavons, and anthocyanins classes) were
reported. The total polyphenol content of the fig fruits was ranged from 715 (‘Verdal’
brevas) to 186 mg 100 g−1 dry matter (‘Campera’). Betulinic acid (content ranging
from 57 to 97%) and oleanolic acid were the main triterpenoids of fig fruits. The
cultivar ‘Verdal’ showed the highest antioxidant activity. The ‘Tiberio’, ‘Campera’,
‘Calabacita’, and ‘Cuello Dama Blanca’ cultivars had significant antidiabetic effect
(Wojdyło et al., 2016).
Extraction conditions (the solvent type and the temperature) and the sample type
affect the quality and quantity of bioactive compounds extracted from fruits. Sun
et al. (2015) isolated phenolic compounds from fresh peeled figs with acidified etha-
nol at room temperature. The combination of alcohol-water showed a better perfor-
mance than mono-component solvents to isolate phenolic compounds. Ethanol and
water are safe and suitable solvents for the food industry than other organic solvents
(Sun et al., 2015). In this regard, Bucic-Kojic et al. (2011) evaluated the effect of
extraction conditions (50–80%, v/v aqueous-ethanol and temperature of 25–80 °C)
and fig variety on phenolic compounds of lyophilized fig fruits. The content of phe-
nolic compounds (2.4–3.7 mg GAE g−1 dw), total flavonoids (0.44–2.5 mg CE g−1
dw), and total proanthocyanidins (0.68–0.87 mg g−1 dw) increased by increasing the
ethanol volume in the solvent from 50 to 80%. The phenolic compound content
increased by increasing temperature (25 to 80 °C). The total phenolic compounds
content in fig fruit was 2.5 to 3.7 mg GAE g−1 dw, while the total flavonoids and
proanthocyanidins content were 0.68–2.5 mg CE g−1 dw and 0.67–0.87 mg g−1 dw,
respectively. Therefore, the best extraction conditions were introduced 80% v/v
aqueous ethanol at 80 °C for 120 min. The highest level of phenolic compounds was
obtained in a variety of Crnica (4.7 mg GAE g−1 dw), followed by Brežutka bijela
(3.9 mg GAE g−1 dw), Šaraguja (3.7 mg GAE g−1dw), Termenjača (3.1 mg GAE g−1
dw), and Bjelica (2.6 mg GAE g−1 dw) varieties (Bucic-Kojic et al., 2011). Meziant
et al. (2014) optimize the extraction conditions (acetone concentration (40–80%),
temperature (25–65 °C), and time (60–120 min) of the total phenolic compounds of
fresh dark fig (Ficus carica L.) by response surface methodology (RSM: The Box-
Behnken design). The optimal extraction parameters were 63.4% acetone,
115.1 min, and 48.66 °C. In this condition, the total phenolic was 536.4 mg GAE
100 g−1, and antioxidant activity was obtained at 71.8 mg GAE 100 g−1 dw (Meziant
et al., 2014). However, organic solvents (e.g., acetone, methanol, ethyl acetate, and
hexane) are related to concerns about environmental pollution, toxicology, and
safety. Developing methods to improve the extraction yield and efficiency of pheno-
lic compounds based on eco-friendly solvents is important. Some papers have
reported that the use of fermentation and enzymatic processes can enhance the
34 The Potential of Fig (Ficus carica) for New Products 775

performance of bioactive compounds extraction from agro-industrial by-products

(Azabou et al., 2016; Larios-Cruz et al., 2019; Torres-León et al., 2021). Fermentation
can enhance the level of bioactive compounds in fruits and vegetables (Martins
et al., 2013). Buenrostro-Figueroa et al. (2017) optimized the solid-state fermenta-
tion (SSF) of fig (Ficus carica L.) by-products by four fungal strains to enhance the
release of total phenolic compounds. The optimized SSF conditions were as fol-
lows: A. niger HT4 for 36 h at 40 °C; moisture content: 60%; pH: 5.0; mineral
composition (g L−1): NaNO3: 0.57; KH2PO4: 3.04; MgSO4.7H2O: 1.52; and KCl:
5.37. Under the optimal conditions, the total phenolic compound was 10.1 mg of
GAE g−1 of dm, and the TPC release and antioxidant activity values increased up to
5.48 and 98.5 times, respectively (Buenrostro-Figueroa et al., 2017). Overall,
quercetin-3-O-rutioside, polymeric procyanidins, quercetin-3-glucoside, chloro-
genic acid, and cyanidin-3-O-rutinoside were introduced as the major individual
phenolic compound in whole figs, while picatechin and cyanidin-3-rutinoside were
the main phenolic compounds of fig pulp (Teruel-Andreu et al., 2021).

3.2 Fig Seed Oil

Along with the progress in the vegetable oils industry, there is a great interest in
using new and cost-effective oil sources. The fig seed oil is healthy. Because it con-
tains ω-3 and rich in unsaturated fatty acids is (Nakilcioğlu-Taş, 2019). The
α-linolenic acid of the fig seed oil is a β-sitosterol and an important omega source
(Güven et al., 2019). Therefore, in recent years, several studies have focused on the
extraction of fig seed oil as a cost-effective source of vegetable oil and evaluated its
quality properties. In this regard, Chougui et al. (2013) evaluated the fig seed com-
position of four varieties of Opuntia ficus-indica. Fig seeds were ground into a fine
powder, and their oil was extracted. The oil content of various varieties varied and
ranged from 7.3% (from the red variety) to 9.3% (from the yellow one). In all the
samples, linoleic acid was the dominating fatty acid (58.7 (red variety) to 63.1%
(orange variety), followed by oleic (15.2% (orange) to 24.3% (red)), palmitic acid
(12.7% (red) to 13.4% (orange)) and stearic (3.3% (orange) to 4.2% (red)). Vaccenic
acid was also only found in the green and orange ones. Gas chromatography analy-
sis of the defatted fig seed powder showed more than 20 phenolic compounds
detected in the defatted fig seed extract (Chougui et al., 2013).
Duman et al. (2018) reported that the oil content of fig seed was 29.1% (acidity:
4.6 mg KOH g−1; iodine value: 176 I2 g100g−1; peroxide level: 4.5 meq O2 kg−1;
refractive index: 1.480 (nD); density: 0.925 mg mL−1; viscosity: 52.1 mPa; melting
point: −15.1 °C; total phenol: 21.7 mg GAE g−1 and oxidative stability: 110 °C h−1).
Linolenic (41.8%), linoleic (29.0%), oleic (18.6%), and Palmitic (6.76%) acids
were reported as major fatty acids of fig seeds oil. Moreover, their results showed
that the fig seed oil had a suitable antimicrobial effect against Staphylococcus
aureus, Pseudomonas aeruginosa, Escherichia coli, Enterococcus faecalis, and
klebsiella pneumoniae bacterial strains besides Candida albicans and Aspergillus
776 S. Khoshnoudi-Nia et al.

flavus as fungal strains (Duman et al., 2018). Hssaini et al. (2020) evaluated the
effect of fig cultivars on fig seed oil. They extracted fig seed oil from four fig culti-
vars. They reported that fig seeds contained 21.5 to 28.5% yellow-colored oil. Gas-
liquid chromatography analysis of the seed oils showed high percentages of linolenic
acid (38.4–43.5) and linoleic acid (28.9–34.5%). The frequent saturated fatty acids
in all fig seed oils were Palmitic acid (8.54 to 9.05%) and stearic acid (2.59 to 3.3%)
(Hssaini et al., 2020). Also, in another work, the total oil yield was 14.08%, and the
antioxidant capacity was 140.1 mg Trolox, equivalent to 100 g−1. The highest fatty
acid was α-linolenic acid (26.3%), followed by linoleic acid (24.2%) and oleic acid
(19.6%) (Ergun & Bozkurt, 2020).
Since fig oil is rich in polyunsaturated fatty acids, it is sensitive to oil oxidation.
Therefore, for more protection, Icyer et al. (2017) encapsulated fig seed oil in gum
arabic (GA) and maltodextrin (MD) as wall material and optimized the encapsula-
tion condition. According to the results, the optimum encapsulation ratio of MD:
GA: oil and spray-drying temperatures were 8:1:3 and 150 °C, respectively (Icyer
et al., 2017).

3.3 Fig Powder

Low-quality fig fruit can be grounded to produce fig powder. The processing tech-
nology used for manufacturing fig powder was simple, and the powder can be used
to develop various functional and value-added food products. Khapre et al. (2015)
prepared the fig (Ficus carica L.) fruits powder (Deanna variety) and used it in the
formulation of burfi (Indian cookie). Fig powder enhanced the nutritional value of
burfi in terms of fiber (3.7%), potassium (0.46%), and protein (13.1%). In addition,
the fig burfi had good taste and low cost compared to similar products (Khapre
et al., 2015). Viuda-Martos et al.(2015) determined the chemical, physicochemical
and techno-functional properties of the fig powder co-products obtained from the
pulp and peel of two cultivars (colar and cuello de dama). The results showed that
the main sugars were fructose and glucose. The sugar, organic acid, and phenolic
acid content of fig powder obtained from the pulp were higher than from fig peel.
However, the flavonoids were predominant in fig powder obtained from fig peel,
and this powder showed higher antioxidant activity than that obtained from the pulp
(Viuda-Martos et al., 2015).
There are many edible seeds in a fig which are a rich source of protein, fat, car-
bohydrates, dietary fiber, antioxidants, phenolic compounds, omega-3 (α-linolenic
and cis- 5, 8, 11, 14, 17-eicosapentaenoic acid), and omega-6 (linoleic acid) fatty
acids and minerals (e.g., Mg, Zn, and Cu) (Baygeldi et al. 2021; Hssaini et al.,
2020). The fig seeds are valuable health-promoting food ingredients. However, dur-
ing the production of various fig products, these seeds are discarded. These wastes
have a good potential to produce value-added products. In this regard, Bölek (2021)
enriched the biscuits with fig seed powder and evaluated its effect on quality param-
eters (proximate composition, color values, total phenolic contents, antioxidant
34 The Potential of Fig (Ficus carica) for New Products 777

activity, texture, and sensory properties) of biscuits. He replaces wheat flour with fig
seed powder at four levels (0, 5, 10, and 15%). Fig seed powder contains suitable
content of fiber (56.6%), protein (14.3%), fat (20.3%), and total phenolic (TPC:
665.1 mg GAE 100 g-1). The addition of this powder to the biscuit formulation
improved the fiber content of the sample and increased the total phenolic com-
pounds and antioxidant activity of biscuits significantly. The addition of fig seed
powder to the biscuit formulation caused an increase in the water absorption capac-
ity, dough stability, hardness, and cohesiveness of the biscuit dough due to the high
content of dietary fiber. The increased dough stability can be referred to their
H-bonding capability obtained by amino acids (especially glutamine and proline) of
fig seeds.
Moreover, the interaction between fiber, fat, and gluten, besides the lower gluten
and higher water holding capacity, leads to a decrease in the gas retention capacity
and spread ratio of biscuits (Kohajdová et al., 2011; Seevaratnam et al., 2012). The
addition of fig seed powder to the formulation of up to 10% enhanced the biscuits’
sensory scores (odors, flavor, and overall score). However, a sensory score of
appearance and texture decreases with the increasing substation ratio of fig seed
powder due to the dilution of gluten caused by wheat flour replacement with fig seed
powder (Bölek, 2021).

3.4 Food Colorant

Food colorants or pigments are natural or synthetic substances used to create,

recover, and standardize food color. Food colorants make food more attractive to
consumers. Anthocyanins are one of the most important water-soluble pigments
with antioxidant activity. They are responsible for the attractive colors of various
fruits and flowers (Yuntao Wang et al., 2017). Therefore, anthocyanins are ideal
alternatives for synthetic dyes with antioxidant activity, which can easily incorpo-
rate into aqueous-based food (Sharif et al., 2020). In addition, the antioxidant activ-
ity of anthocyanin induces various health benefits in preventing cancer (Santos
et al., 2013), neuronal and cardiovascular illnesses (He & Giusti, 2010), inflamma-
tion (Lee et al., 2011), and other diseases.
Peel of fig has a significant potential to isolate phenolic compounds, especially
anthocyanins (Buenrostro-Figueroa et al., 2017). In this regard, a high concentra-
tion of phenolic compounds and anthocyanins also were reported in the peel of
mature figs (Barolo et al., 2014). Backes et al. (2020) used natural purple colorants
extracted from fig peel in doughnuts (icing) and a typical Brazilian pastry called
“beijinho” formulation. The extract of fig peel had no significant changes in nutri-
tion value and rheological features. Even the dough firmness and consistency were
improved, and “beijinhos” became softer and chewier. Moreover, the extract showed
antioxidant and antimicrobial activities in the products (Backes et al., 2020).
778 S. Khoshnoudi-Nia et al.

3.5 Fig Pectin

On an industrial scale, Pectin, an abundant structural acidic polysaccharide, is com-

monly produced from citrus, beet, and apple wastes. Due to gelling and emulsifying
properties of pectin, it is an excellent candidate for stabilizing jams, jellies, and
acidic dairy drinks (Brejnholt, 2009). Moreover, it can also be used to formulate
micro-and nano-capsules and edible films/coating (Kurek et al., 2021). In addition,
various health benefits were introduced for pectin, such as decreasing cholesterol
and serum glucose levels, preventing cancer, and improving the immune system
(Wikiera et al., 2014). Therefore, there is a growing interest in isolating this valu-
able and multifunctional polysaccharide from new and non-expensive sources,
especially fruit peels and pomace. In this regard, Gharibzahedi et al. (2019) used
four extraction methods (i.e., hot-water, ultrasound-assisted, microwave-assisted,
and ultrasound-microwave assisted) to extract pectin fig (Ficus carica L.) peel. The
most yield was obtained for ultrasound-assisted extraction (11.7%). The fig peel
pectin, especially the sample extracted by the ultrasound-assisted, exhibits a rela-
tively good antioxidant activity-dependent. Moreover, the pectin extracted by the
ultrasound-assisted method presented the maximum galacturonic acid content
(76.8%), molecular weight (6.91 × 103 kDa), the highest emulsifying activity
(61.2–61.3%), and emulsion stability (94.3–95.2%) (Gharibzahedi et al., 2019).

4 Conclusion

Nowadays, there is an increased interest in developing novel nutrient-rich food

products. There is scientific research about the potential use of fresh, dried, and
underutilized fig fruit and figs by-products to produce new and value-added food
products. The use of low-quality figs and their waste to produce food ingredients
and bioactive compounds could benefit the food industry, reduce environmental
issues, and benefit consumers’ health. Although fig-based products were reported,
the effect of novel technologies for extraction of fig bioactive compounds and using
these valuable extracted compounds in food formulation to improve the quality,
shelf-life, and nutritional value of food products needs further study. In addition, the
use of fig waste to produce novel products such as fig coffee and fermented fig milk
should be investigated.

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Chapter 35
Fig Production and Processing: A Pakistan
Perspective

Aijaz Hussain Soomro and Tahseen Fatima Miano

Abbreviations

AJK Azad Jammu and Kashmir

ha Hectare
KP Khyber Pakhtunkhwa
USA United States of America

1 Introduction

Nature has bestowed Pakistan with a wide range of agro-ecological regions such as
tropical, subtropical, and temperate regions and thus a diversification in growing
many horticultural produces (Ahmad et al., 2021). Among different horticultural
crops, the fruits grown in Pakistan have considered highly economical and impor-
tant crops for potential exportation, local consumption in raw or processed form,
and food insecurity (Shahzad et al., 2019). Moreover, fruits are the oldest form of
natural staple food for humanity (Khadivi et al., 2018; Alam et al., 2021).
Fig is one such fruit that is supposed to be the oldest fruit domesticated in
Mediterranean regions (Kharadi et al., 2005) and is native to western Asia and the
eastern parts of Mediterranean countries (Hmimsa et al., 2012). It is believed that
the fig fruit was domesticated 5000 years earlier than wheat and millet (Khan et al.,
2022). The consumption of fig fruit dates back to the beginning of civilization
(Ferraz et al., 2021). This fruit is valuable agricultural produce grown in tropical
and subtropical regions worldwide. Since ancient times, the fig has been a part of
the diet and is considered the symbol of heaven and longevity. It is a popular fruit

A. H. Soomro (*) · T. F. Miano

Institute of Food Sciences and Technology, Sindh Agriculture University,
Tandojam, Sindh, Pakistan
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 785
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_35
786 A. H. Soomro and T. F. Miano

worldwide and is known by different names in different languages. The fruit is

named Teen in Arabic, Anjeer in Urdu, Gujrati, Hindi, Kashmiri, Figu in French,
and Anjir in Turkish (Nuri & Shahabuddin, 2021). There are 470 varieties of edible
fig. Commercially grown varieties are Celeste, Brown Turkey, Brunswick,
Marseilles, Adriatic, Genoa, Purple Genca, and Black Ischia. These varieties are
grown in different regions of the world. Among them, Black Mission and Brown
Turkey are highly produced varieties. In Pakistan, the fig cultivation was 162 ha,
with around 487 tons. The average fig production is about 3781 Kg per ha. This
indicates that the yield of fig fruit in Pakistan is very low compared to other coun-
tries. Despite having suitable agro-climatic conditions in the prospect of fig fruit
production, Pakistan is far enough due to the lack of interest of growers and other
stakeholders. However, countries like Turkey, Egypt, and Iran are the leading pro-
ducer of fig fruit globally. The fig fruit is also cultivated in Algeria, Morocco, Syria,
the USA, Brazil, Spain, India, Albania, and Tunisia (Arpaci et al., 2018).

2 Fig Tree and Its Cultivation

Fig plant is a member of the Moraceae family, mainly cultivated for its valuable
fruit (Kim et al., 2007). Fig tree bears various horticultural properties, and it offers
fresh fig fruit twice a year during the spring and summer seasons (Aksoy, 2017).
This tree may ideally grow in rural areas; however, it is badly neglected and under-
utilized, especially around Mediterranean regions from where it originated. The fig
trees are mainly cultivated in Pakistan’s KP, Punjab, and Sindh Provinces. In Azad
Jammu and Kashmir (AJK), the fig fruits are grown in the natural forests, and mar-
ginal arable land with lower economic returns since the fruit is mainly consumed
locally (Qureshi et al., 2007). However, the climatic condition in Pakistan provides
an ultimate opportunity for the growth and development of fig trees. Different fig
varieties, namely English fig, Wild fig, Dark brown, and Dark black figs, are among
some common fig varieties in Haramosh Valley, Gilgit, Pakistan (Abbas et al.,
2016). The common fig varieties grown in different countries are presented in
Table 35.1.
The fig tree is a deciduous shrub or a small tree that is mainly adaptive to differ-
ent soil types and climatic conditions and is widely grown in many parts of the
world. The tree grows fast with a spreading habit, the flowers of the fig tree are tiny
enough, and the fruit is hollow (Stover et al., 2007). The seeds of fig fruit are non-
viable; therefore, fig trees are propagated via cutting or grafting. A fig tree needs
sunlight for around eight hours a day for proper fruit ripening; however, fruits must
be allowed on the tree for ripening. The nutrient uptake of the fig tree increases dur-
ing the developmental stage of fig fruit. Fig trees occur in bisexual forms, i.e., a
functional male caprifig and other unisexual females which produces edible fig fruit
(Kim et al., 2007).
35 Fig Production and Processing: A Pakistan Perspective 787

Table 35.1 Fig varieties and their regions of cultivation

Fig varieties Region of cultivation Reference (s)
Wild Himalayan fig AJK, Pakistan Khan et al. (2022)
Green, Purple, and Black figs Tunisia Harzallah et al.
Mediterranean coast (2016)
Mission, Vasilika, and Mavra Markopoulou figs Southern Europe Karantzi et al.
(2021)
Sarilop, Bursa Siyahi, Goklop, Yediveren, Yesilguz, Dortyol, Hatay, Çalişkan and Polat
Morguz, Sari Zeybek, and Ufak Yesil figs Turkey (2008)
Khafrak, Rowno, Sigoto, Shahanjir. Seyah, Matti Fars province, Iran Pourghayoumi et al.
figs (2016)
Conadria, Deanna, Excel, Poona, Dinkar figs India Gawade and Waskar
(2005)
Roxo de Valinhos, White Genova, PI-189, Troyano Brazil Ferraz et al. (2021)
figs
Mission, Chechick, Bursa, Brown-Turkey, Bet-Dagan, Israel Solomon et al.
Brunswick, Kadota figs (2006)
Aberkane, Azandjar, Bouankik figs Algeria Bachir Bey et al.
(2017)

3 The Scenario of Fig Production in Pakistan

In Pakistan, the fig trees are mainly cultivated in KP, Punjab, and Sindh Provinces.
In Azad Jammu and Kashmir (AJK), the fig fruits are grown in the natural forests
and marginal arable land, with lower economic returns since the fruit is mainly
consumed locally. The total area under fig cultivation in Pakistan is about 125 hect-
ares which is very low compared to the rest of the world (Qureshi et al., 2007).
However, it is factual that the climatic condition in Pakistan provides an ultimate
opportunity for the proper growth and development of fig trees. Different fig variet-
ies, namely English fig, Wild fig, Dark brown, and Dark black figs, are among some
common fig varieties in Haramosh Valley, Gilgit, Pakistan (Abbas et al., 2016),
while varieties like Black Turkey and Black Mission are also two prominent variet-
ies of fig grown in Pakistan (Amjal et al., 2016).

4 Fig Fruit

Fig fruit is an edible part of the fig tree which is pear-shaped, fleshy, receptive, hol-
low fruit (Dueñas et al., 2008), sweet with gelatinous pulp (Stover et al., 2007) is
termed as syconium. This fruit may be consumed in fresh or dried form while both
forms are nutritionally superior and deliciously palatable. The developmental stages
of growing fig fruits are different from the other fruits (Heperkan et al., 2012).
Usually, the fruit may vary in size and color depending on the variety (Farhangi
et al., 2014). The fruit size may range from 3 to 5 cm, while a mature fig fruit has a
788 A. H. Soomro and T. F. Miano

tough brown skin that often cracks when the fruit ripens/softens and exposes the
fruit pulp in beneath. The color of figs varies from dark purple to green. The fig fruit
varieties with dark skin color are rich in polyphenols, i.e., flavonoids and anthocya-
nins (Solomon et al., 2006). This fleshy fruit bears many subtle and crunchy seeds
and is embedded in the fruit pulp so that the seed mass is bound with the jelly-like
thick material within the inner skin of fig fruit. The pulp of fig fruit is mainly water,
minerals, volatile compounds, and vitamins, while the seeds have a nutty taste. The
skin of fig fruit is loaded with many polyphenols and exerts antioxidant activity in
darker fig varieties than lighter ones (Solomon et al., 2006). In addition, the fruit
contains volatile components providing a pleasant and characteristic aroma (Bachir
Bey et al., 2017).

5 Nutritional Composition of Fig Fruit

Fruits from the fig tree are a true gift from nature. The fruit contains many essential
components associated with maintaining health and wellbeing (Soni et al., 2014).
The fruit bears significant varietal differences. Thereby composition of fruit varies
significantly amongst cultivars. The fig fruit bears medicinal properties, has an
excellent nutritional profile, and is low in calories, i.e., 74 calories/100 g of fresh
fruit (Farhangi et al., 2014). Both fresh and dried fig fruits are rich in fiber, minerals,
organic acids, and antioxidant vitamins (A, E, and K) while are free from choles-
terol (Stover et al., 2007). Fruit is also the richest source of amino acids
(Bharathkumar et al., 2018), particularly aspartic acid and glutamine. The fruit is
loaded with minerals and is considered a healthy fruit since it is rich in many essen-
tial minerals, i.e., iron, potassium, calcium, magnesium, sodium, and zinc (Sadia
et al., 2014). Fresh fig fruit has an average dietary fiber and protein content while
very low-fat content (Soni et al., 2014). According to Vinson et al. (2005), fresh and
dried figs contain more fiber and polyphenols. Solomon et al. (2006) also reported
that the higher the polyphenols contents, especially anthocyanins in fig fruit, the
higher was their antioxidant activity. Figs also comprised sugars and organic acids
that influence their quality and exhibit one of the highest concentrations of polyphe-
nols among the commonly consumed fruits and beverages that contribute positively
to human health (Veberic et al., 2008). Mahmoudi et al. (2018) found that the peels
and pulps of figs have good nutritional value. The fruit pulps are rich in carbohy-
drates, and the peel of the ‘Onk Elhamam’ cultivar had the highest vitamin C con-
tent. The pulp of the ‘Boughandjo’ cultivar contained the highest concentration of
phosphorus, and the peel of the ‘Bakkor Khal’ cultivar was richest in potassium and
calcium.
Moreover, Ersoy et al. (2007) suggested that fig fruit contains sugars such as
maltose, xylose ribose, arabinose,e, etc. Fig fruit is a good source of health-
promoting bioactive components (Harzallah et al., 2016). However, dried fig fruit
contains a relatively higher concentration of crude fibers (approximately 5.8%),
sugars, and polyphenols (Vinson et al., 2005). It also has a reasonable proportion of
35 Fig Production and Processing: A Pakistan Perspective 789

B-complex vitamins such as niacin, pyridoxine, pantothenic acid, folates (Farhangi

et al., 2014), and minerals, i.e., potassium, and calcium, magnesium, and sodium
(Turco et al., 2020). In Azad Jammu and Kashmir, edible figs are generally grown
in natural forests and marginal lands, with very low economic returns as most of the
fruit is consumed Since figs are perishable, they are dried naturally under sunlight
in Kashmir. Figs produced in central Kashmir have a very high fiber content. They
are also reported to be an excellent source of calcium and potassium. Dried Kashmiri
figs contain omega-3 and omega-6 fatty acids, which are considered helpful in pre-
venting heart disease and can cure sore throats due to their high mucilage content
(Mir et al., 2018).

6 Medicinal Properties of Fig Fruit

The consumption of figs has positive health effects due to the numerous nutraceuti-
cal compounds that may help prevent cardiovascular diseases and the growth of
carcinoma cells (Allegra et al., 2017). Some fig varieties contain many therapeutic
compounds, thereby helping traditional ethnomedical remedies. Almost all parts of
its tree (leaf, fruit, latex, root, etc.) have medicinal effects against innumerable dis-
eases. Many species of Ficus carica are potent sources of antioxidants with strong
anti-microbial, anti-diabetic, anti-inflammatory, anti-spasmodic, and anti-
obesogenic activities (Shahrajabian et al., 2021). The fig tree's fruit plays an essen-
tial role against numerous health disorders related to oxidative stress (Ahmad et al.,
2013). Thereby fig fruit immensely contributes to good health and wellbeing. The
phytochemical components in the fruit scavenge toxic free radicals from the body
and exhibit protection against cancer, diabetes, and infectious and degenerative dis-
eases (Farhangi et al., 2014). According to Shahrajabian et al. (2021), dark-skinned
fig varieties are rich sources of polyphenols, flavonoids, anthocyanins, and pro-
anthocyanidins in exerting antioxidant activity than light-skinned fruit. Steam distil-
lation of fig fruit contains benzaldehyde which exerts considerable antitumor action.
Interestingly, Solomon et al. (2006) suggested that fig fruit is a useful mild laxa-
tive, diuretic, and expectorant in Indian Ayurveda. At the same time, its paste is
often used against swellings and for relieving pain. Fresh fig is valued for its laxa-
tive effects, treating skin infections, and maintaining acid-base balance in the body.
Apart from the fig fruit, the other parts of the fig tree (roots, leaves, etc.) are report-
edly used as traditional medicines and are immensely effective against specific car-
diovascular, gastrointestinal, and respiratory ailments (Khan et al., 2022). Moreover,
fig leaves' decoction (a method of extracting a plant’s medicinal constituents by
boiling the plant material) is effective against ameliorating post-prandial hypergly-
cemia. A reduction in hypercholesterolemia and hyperglycemia has been further
utilized to treat the aqueous decoction of fig tree leaves.
On the other hand, the fig leave extract may induce a significant hypoglycemic
effect after oral or intraperitoneal administration. Patients with Diabetic Mellitus
may benefit from fig tree leaves; therefore, researchers should explore its influential
790 A. H. Soomro and T. F. Miano

role in Diabetic Mellitus (Ajmal et al., 2016). Many pathogenic bacteria pose health
problems because of the multidrug resistance that may have developed due to the
antibiotics used against infections. The fig possesses antimicrobial properties
against some pathogenic microorganisms. For example, the dried fig extracts inhib-
ited the growth of Bacillus subtilis and Proteus mirabilis (Soni et al., 2014).

7 Perishability of Fig Fruit

The fig fruit has tremendous commercial importance (Irget et al., 2008), but unluck-
ily the fruit is highly perishable and has a scanty shelf-life. The major reasons for its
perishable nature are fungal attack, weight loss, mechanical damage, etc. (Colelli &
Amodio, 2020). It has a short shelf-life compared to other fruit due to its high mois-
ture content and sugar concentration. Fresh fig fruit's moisture content may range
from 70.5% to 82.3% (Bachir Bey et al., 2017). In its new form, the fruit is highly
vulnerable to microbial attack even under the cold-store facility or refrigeration
conditions (Farahnaky et al., 2009). Due to its immense perishable nature, the fruit
is consumed in its fresh raw form or dried. A fully ripened fresh fig fruit has a shelf-
life of 2–3 days under refrigeration conditions. Fruit takes about 4–5 days for sun
drying while 10–12 h under dehydrator. The United Nations Economic Commission
for Europe-Standard (2004) suggests that moisture content in commercial-grade
dried fig fruit should not exceed 26%. The dried figs have prolonged storability of
about 6–8 months.

8 Fungal Invasion and Mycotoxin Production in Fig Fruit

Fig is an economically important fruit crop and is susceptible to fungal infection.

Fig fruits are at high risk of toxigenic fungi. The fungal invasion may occur in the
fig tree after fruit shriveling or falls and even during their drying. The fungal growth
is often associated with the subsequent production of mycotoxin (Heperkan et al.,
2012). The water activity of fig fruits during their production or processing affects
the growth of toxigenic fungi and the formation of mycotoxins. The climatic condi-
tions like high humidity %, mild temperatures, and rain promote fungal growth. The
dominant fungal species that occur in dried figs may vary widely, and their types
depend on different factors such as geographical regions, processing methods, etc.
Some common types of reported toxigenic fungi are Aspergillus spp., Fusarium
spp., and Penicillium spp. Aflatoxins and ochratoxins occur more widely in fig fruit
(Heperkan, 2006).
35 Fig Production and Processing: A Pakistan Perspective 791

9 Fig Fruit Processing and Value Addition in Pakistan

The perishable nature of fig fruit tends to fruit growers and processors for its suit-
able processing and effective value-addition. A main proportion of the fruit is sub-
jected to drying or dehydration to develop dried figs. The processing technologies
are employed on fig fruit to attain more imperative and valuable commercial fruit-
based commodities. However, a significant number of scientific studies have been
conducted by many researchers for developing fig fruit-based food products
(Table 35.2). Pakistan lacks suitable fig processing industries since the fruit is pro-
duced or cultivated in marginal quantities. The fruit's industrial processing and tech-
nological aspects may attain a prime place in the country’s economy if serious
measures for a sustainable fig production system are taken. It is worthful that
increased demand for fig fruit (fresh or processed) worldwide is gaining efforts to
expand its production; therefore, Pakistan is also paying keen attention to increased
cultivation of figs in different arable areas of the country. However, it is also impera-
tive for the food processing industries in Pakistan to mitigate the post-harvest losses
of figs by applying technological practices during processing and preservation with
improved drying techniques and preparation of fig-based value-added products.

10 Fig-Based Value-Added Food Products

Efforts are needed to increase the production of fig fruits to meet market demand in
Pakistan since the production of the commodity is associated with its processing.
However, the comprehensive research study for exploiting the diversity in wild figs

Table 35.2 Utilization of fig fruit for the development of value-added food products
Fig fruit form Use in value-addition References
Fresh fig fruit Fig jam and preserve Levaj et al. (2010)
Fig jam and nectar Tanwar et al. (2014)
Dried fig Slatnar et al. (2011)
Novel functional fig snack Yeganehzad et al. (2020)
Canned figs Curi et al. (2019)
Fig wine Jeong et al. (2005)
Fig fruit-based fermented milk Abd-Eltawab and Ebid (2019)
Dried fig Fig powder Khapre et al. (2011)
Fig acnes/seeds oil Soltana et al. (2016)
Hard candy Verma and Gupta (2015)
Fig wine Kadam et al. (2011)
Fig powder Toffee Khapre et al. (2011)
Cookies Khapre et al. (2015a)
Burfi (Indian cookie) Khapre et al. (2015b)
Fig paste Fat replacer in baked foods and sauces Benjamin (2020)
792 A. H. Soomro and T. F. Miano

is not conducted for their possible use in sustainable fruit production. Subsequently,
propagation and production of the species in the region may contribute to upgrading
the local economy and may significantly influence socio-economic and ecological
stability. Therefore, the local farmers have good opportunities for producing wild
figs to increase income generation and provide a sustainable natural food source to
the consumers (Khan et al., 2022). However, the perishable nature of the fig fruit
also causes significant loss to the country’s economy.
Moreover, it is also direly essential to reduce its waste (or utilize surpluses) by
preventing or controlling the post-harvest losses of the fruit. The fruit thereby needs
to receive proper preservation/ processing techniques that should be simple and eas-
ily adaptable. Benjamin (2020) suggested that fig-based ingredients of different
grades have been established to have attributes like natural coloring and flavoring
agents, humectants, and anti-staling agents. Fig fruits may be used to develop many
value-added food products. Sher et al. (2016) suggest that presently, the fruit is
being utilized for producing various food commodities (jams and jellies, etc.). In
addition, the fig fruit may be utilized as an optional ingredient in baked items (cakes,
bread, biscuits, and pies) and for garnishing.

10.1 Dried Fig

Drying is the most popular and effective way of processing/preserving figs known
from prehistoric times (Kislev et al., 2006). Drying and dehydration have proven to
be reliable preservation techniques for fig fruit in terms of nutritional quality and
technical feasibility (Desa et al., 2019). Drying has many advantages for food qual-
ity decreasing water activity, reducing microbiological activity, and minimizing
physical and chemical changes (Mujic et al., 2014). During fig drying, the
temperature-time relationship plays a crucial role. The global market of dried fig
fruit is expanding; however market always demands safe, hygienic, and high-quality
dried figs. Dried figs comprise about 90% of the world's production. Dried figs are
distinguished by high nutritional value and possess functional food properties
(Vinson, 1999). Galván et al. (2021) suggest that the processing of dried fig fruit has
increased by 44% in the last decade. The high energy demanding artificial drying
methods are unreliable and expensive, while in contrast, slower methods like sun
drying or solar drying are suitable alternatives. Therefore, natural sun drying is a
widely practiced drying method in tropical and subtropical regions. During their
drying, storage, and retailing, dried fig processing should be considered since fruits
may be susceptible to fungal attacks or mycotoxin production (Devreese et al.,
2013). The suggested ambient temperatures, i.e., 16 and 25 °C, may be considered
reliable against fungal growth for both processors and consumers for storing dried
figs (Galván et al., 2021). Dried figs are a good source of carbohydrates, sugars,
minerals, vitamins, organic acids, and phenolic compounds (Veberic et al., 2008).
Both fresh and dried figs have high fiber and polyphenols (Vinson, 1999; Vinson
et al., 2005).
35 Fig Production and Processing: A Pakistan Perspective 793

10.2 Fig Powder

The technology for producing fruit powder is gaining significant importance. Fig
fruit is generally subjected to drying and then milled to obtain fig powder. The fig
powder is considered ready-to-eat food (Varhan et al., 2019), whereas it may also be
used as a food ingredient. Varhan et al. (2019) suggested that drying technologies
preferred to produce food powder are spray-drying, freeze-drying, tray-drying,
foam-mat-drying, microwave drying, etc. Fig powder is nutritionally rich and con-
tains reasonable quantities of proteins, fats, minerals, and sugar (Khapre et al.,
2011). Nowadays, there is an increase in nutrient-rich value-added products' devel-
opment by partially replacing its ingredients with others, such as underutilized
fruits and added value by-products (pectins, colorants, emulsifiers, and antioxi-
dants) from leaves and peels. As for fig by-products, the use of fig powder as a colo-
rant in the production of buns and muffins (Chauhan & Tanwar, 2016). The addition
of fig seed powder to the formulation of a cookie also improved its fiber content and
increased the total phenolic content and antioxidant activity (Bolek, 2020). Fig fruit
cake is a widely used snack by children and adults (Dhankar, 2013).
Additionally, fig by-products’ sweet extracts have been used for making tradi-
tional desserts without adding sugar, for example, “Shir Anjir”, an Iranian dessert
(Jahromi & Niakousari, 2018). The fig powder supplement, used as a natural sweet-
ener and flavoring agent in goat’s milk yogurt, has improved the taste, texture, and
aroma and covered the unpleasant flavor of goat’s milk (Mahmoudi et al., 2021). Fig
powder may be utilized for making various food products (Table 35.2).

10.3 Fig Jam

Fig jam making is a popular approach for preserving fig fruit. The fig jam prepared
from the freshly harvested and fully ripened fig fruit is a rich source of nutrients.
The fig jam is a palatable and nutritious food item and has tremendous market
potential. The figs are usually available commercially after drying because fresh
fruits are only available during the season. The fresh fruits have a short shelf life,
usually 7–10 days. Therefore, figs could be preserved through jam processing. In
Bulgaria, fig jam with whole green and ripened fruits is a very popular type of pres-
ervation of these seasonal fruits. (Rababah et al., 2011; Tanwar et al., 2014).

10.4 Fig Paste

Dried figs may be utilized to produce versatile food products, for instance, fig con-
centrate, fig butter, and fig paste. Fig paste is a slurry that may be incorporated in
many food formulations. The quality of fig paste may vary depending upon the
794 A. H. Soomro and T. F. Miano

variety of fig, its particle size, and the final moisture content of the paste slurry
(Benjamin, 2020). The fig paste is a versatile ingredient in terms of technical func-
tionalities. Fig paste form colloidal bonds with components of the food matrix (fats,
protein, water, etc.). Fig pastes and water slurry ratio of 50:50 is considered excel-
lent emulsification with suitable sensory appeal and physicochemical properties.
Therefore, fig paste may be used as an ingredient in food products. Traditionally, fig
paste is prepared by selecting mature and fresh fig fruits. After washing, figs are
cooked with the addition of sugar on a slow flame by occasionally stirring for a few
hours till a homogenous slurry of paste is achieved (Lee et al., 2012).

10.5 Fig Seed Oil

Fig seeds may be separated from the fruit, dried, and processed to obtain fig oil. Dry
seeds contain 30% oil (Joseph & Raj, 2011). The fatty acid compositions of the fig
seed contain a significant amount of α-linolenic, linoleic, and oleic acids. In addi-
tion, fig seed also contains a small quantity of palmitic acid. Linolenic and linoleic
acids are essential fatty acids and constitute about 70% of the fig seed fatty acids
(Baygeldi et al., 2021). In addition, omega-3 fatty acid reduces the risk factors for
cardiovascular and age-related macular degeneration due to their anti-inflammatory
effect (Richer et al., 2016).

10.6 Fig Pickle and Fig Chutney

Fig fruit pickle can be prepared by selecting good quality fig fruits using different
food ingredients such as oil, vinegar, sugar, and spices, while fig fruit chutney is
prepared by adding whole fig fruit paste, brown sugar, apple cider vinegar, chopped
onion, and spices.

10.7 Canned Figs

Many forms of perishable fruits are usually canned to prolong their shelf life.
Canning is an old method of preserving fragile food commodities. Fruits with
shorter shelf life may be canned to increase their shelf life. In general, figs are char-
acterized by their high moisture content. Usually, 82% are considered a great source
of carbohydrates of which the principal sugars are soluble, responsible for their
sweet taste, and present high perishability, making their fast transportation to the
centers of consumption necessary. The leading causes of losses in fig quality are
inadequate harvest and packing and the lack of standardization in the classification
of the product (Caetano et al., 2017). The canned fruits prepared from the different
35 Fig Production and Processing: A Pakistan Perspective 795

fig cultivars may have different textures. The amount of sugar present in each culti-
var, pH, and acidity can influence gelling and the final product's texture (Souza
et al., 2014). In addition, aspects such as moisture content and the chemical compo-
sition of the fruit may also change texture, as it may affect cooking time, yield, and,
therefore, the moisture content of the processed product (Curi et al., 2017). Curi
et al. (2019) prepared eight different formulations of canned fig fruits and evaluated
them for their quality attributes. Conclusively, fig fruit supplementation in the diet
has been beneficial to coping with various metabolic disorders due to their high
nutritional potential and antioxidant activity.

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Chapter 36
Wound Healing and Ficus carica (Fig)

Nahla A. Tayyib

Abbreviations

FC Ficus carica
HE Hematoxylin and Eosin
MT Masson’s Trichome
SWA Scratch wound assay
BHK 21 Baby Hamster Kidney
MDCK Maddison-Darby Canine Kidney

1 Healing Powers of Fig

Figs are rich in calcium, iron, and vitamin A in the form of carotene, while niacin
and riboflavin are found in fewer amounts. Three fresh figs contain 140 IU of vita-
min A, 0.5 mg of iron, 15 mg of calcium, 0.2 mg of niacin, and 0.03 mg of ribofla-
vin. Two dried figs contain 20 IU of vitamin A, 1.4 mg of iron, 35 mg of calcium,
0.5 mg of niacin, and 0.04 mg of riboflavin (Ben-Noun et al., 2003).

2
Ficus carica and Its Enzymes

Figs are among the sweetest widely available fruits, a member of the mulberry fam-
ily (Arvaniti et al., 2019). Figs were employed as a sweetener in ancient times, well
before the discovery and production of refined sugar. In the first verses of Sura

N. A. Tayyib (*)
Nursing Practice Department, College of Nursing, Umm Al-Qura University,
Makkah, Saudi Arabia
e-mail: [email protected]

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 801
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4_36
802 N. A. Tayyib

Al-Teen, the Quran discusses the medicinal benefit of the figs: “I swear by the Fig
and the Olive”. In addition to pomegranates, olives, grapes, bananas, and dates, the
Quran mentions the fig plant as a fruit (Asma Arshad, 2014; Mehdi Trad et al. 2014;
Yadav et al., 2015). The fig is the fruit of Ficus carica (FC), a tiny tree in the
Moraceae family of flowering plants (Barolo et al., 2014). In the Mediterranean and
western Asia, it has been growing since antiquity and is widely grown for its fruit
and as a decorative plant worldwide (Mehdi Trad et al. 2014). In addition, the fig
tree is extensively mentioned in the Bible and has existed for thousands of years in
the Middle East and North Africa. Figs are not only delicious but also incredibly
healthy fruit (Dafni & Böck, 2019).
An investigation by Gagaoua et al. (2014) found that figs are one of the few fruits
with an enzyme capable of protein digestion. Moreover, several researchers (Kirtikar
& Basu, 1975; Warrier, 1996; Chopra et al., 1992) argued that Ayurveda had
employed various components of this plant and latex, either raw or in different for-
mulations, to cure wounds. Findings revealed the properties of fig leaf extract, an
enzyme of fig that degrades proteins in enhancing the efficacy of medicines against
infections such as Staphylococcus. Furthermore, ficin, found in the latex of fig, is
one of the most promising proteases developed from plants (Ficus carica), capable
of metabolizing protein into amino acids.

3 Wound Healing Properties of Fig in Eliminating Biofilms

The wound healing properties of figs have generated a broader interest in investigat-
ing the role of the specific parts of the fruits (Harish et al., 2008). Studies have
shown that treatment with ficin reduces the wound‘s half-closure time by two days
and accelerates its microbial decontamination by four days (Baidamshina et al.,
2020). Model wounds of laboratory animals treated with immobilized ficin were
cleansed of microorganisms and healed more quickly. Moreover, the structure of the
newly created tissue treated with protease possesses properties most similar to those
of the original, intact tissue, Kayumov et al. (2015) emphasized. In another study,
Baidamshina et al. (2017) stated that Staphylococcus-formed biofilms frequently
impede the treatment of wounds. A study from the University of Kazan discovered
that ficin, an enzyme from fig latex, is effective for eliminating these biofilms, which
could be a significant barrier to wound care. Within the biofilm, antibiotics cannot
reach the bacterium. Interestingly, increasing the antibiotic concentration by as
much as a thousandfold in wound treatment is one option. Bacteriophages, an anti-
bacterial treatment regaining favor, can function within biofilms.
In an in vitro study, chitosan-immobilized ficin destroyed staphylococcal (S) bio-
films, thereby enhancing the antibacterial activity against biofilm-embedded bacte-
ria. In vivo, in the presence of ficin (either soluble or immobilized), the skin wound
areas of rats infected with S. aureus decreased by twofold after four days as opposed
to six. Moreover, topical treatment of the immobilized enzyme resulted in a 3-log
reduction of S. aureus cell count on wound surfaces in 6 days. In contrast, the
36 Wound Healing and Ficus carica (Fig) 803

control group required more than ten days to achieve the same outcome. Therefore,
additional benefits include smoother re-epithelization and the generation of new
tissue with collagen structural properties that closely resemble those observed in the
native tissue. Both soluble and immobilized ficin appear effective for the treatment
of biofilm-associated infections, as well as for accelerating wound healing and
microbial decontamination (Baidamshina et al., 2020).
Bacterial cells are particularly resistant to antibiotics and immune system
responses in biofilm, resulting in chronic reinfections. In addition, numerous oppor-
tunistic bacteria (Staphylococcus, Micrococcus, Klebsiella, Pseudomonas, etc.)
develop biofilms on chronic and acute cutaneous wounds, preventing their healing
and causing reinfection and sepsis (Worthington et al., 2013). Ficin is a safe and
effective medication for treating external wounds to inhibit biofilm formation and
reduce the risk of reinfection (Baidamshina et al., 2017). Biofilms are a thin but
resilient collection of surface-associated microbial cells encased in an extracellular
matrix, and bacteria are protected from antibiotics within these biofilms. Moreover,
one option in wound care is to increase the antibiotic concentration by as much as a
thousand-fold to push bacteriophages off the ledge in an antimicrobial therapy that
could function within biofilms (Harper et al. 2014).
Numerous studies have been conducted on plants used to hasten wound healing.
According to patents and articles, many herbal compositions expedite wound heal-
ing and could be used to treat wounds. The fig leaf is one of the therapeutic herbs
used (Ficus carica Linn). Fig leaves contain flavonoids, terpenoids, tannins, alka-
loids, and saponins. It is recognized that these chemicals in tin leaves have biologi-
cal action as antioxidants, anticancer agents, anti-inflammatory agents, antiviral
agents, and antibacterial agents. There are hundreds of different types of figs. In Tib
al Nabawi, this fruit is utilized as a medicine in the form of large tablets resembling
crushed fruit (Mutiara Figs, 2010).

4 Ficin and Its Medicinal Profile from Different Parts of Fig

Different parts, particularly the latex, of Ficus racemosa L. have been employed as
a wound-healing medication in Ayurveda and traditional medical systems world-
wide. As a result, there are large populations of Ficus racemosa L. (Family
Moraceae) worldwide, including Asia, Africa, America, and Australia. They are
known as Cluster Figs in English (Yadav et al., 2015). Ayurveda also employs a
variety of raw and prepared forms of this plant’s components and latex in healing
wounds. Medicinal herbs for wound healing and skin illnesses are the key to fight-
ing pathogen resistance to medicines and allopathic treatment. Consistent with
Baidamshina et al. (2017), researchers in Russia have explored the influence of
ficin, an enzyme that degrades proteins, on the efficacy of medicines against
Staphylococcus infections. In Russia, researchers at Kazan Federal University and
Voronezh University have focused on an understudied protease isolated from fig
latex. As reported in scientific reports, the enzyme successfully removed biofilms
from Staphylococcus bacteria in preclinical trials.
804 N. A. Tayyib

Another report (Labiotech, 2017) documented that Staphylococcus bacteria,

which could cause severe infections and are widespread in skin lesions, find it easier
to degrade biofilms due to their comparatively high protein content. Proteases such
as trypsin and papain extracted from papaya were previously utilized in wound care.
However, ficin appears to be more effective than the regularly used trypsin. Plant
proteins, such as papain and ficin, have a low allergenic profile and are typically less
cytotoxic than animal or bacterial enzymes. Murti et al. (2010) highlighted that figs
play a significant role in the treatment or maintenance of wounds. They also men-
tioned that herbal remedies, such as plant-based figs, could significantly enhance
wound disinfection, debridement, and the provision of a moist environment, which
helps the creation of an adequate natural healing climate. The traditional Indian
medical system considers the entire plant’s parts (leaves, fruits, bark, latex, and root
sap) medicinally significant (Murti et al., 2010).
Bopage et al. (2018) evaluated the properties of the trunk bark that is abundant in
sterols, including sitosterol, lupenol, and stigmasterol. In addition, they stated that
the fruits have gluanol acetate, which is the principal constituent. The remaining
components include hydrocarbons, glucanol, tiglic acid, taraxasterol, lupeol ace-
tate, friedelin, and lupeol acetate. The many forms of steroids included in latex
include euphol, isoeuphorbol, sitosterol, 4-deoxyphorbol, cycloartenol, and cycloe-
uphordenol. It was revealed that the hexanes and dichloromethane extract of the
stem bark of F. racemosa have a statistically significant effect on wound healing,
and the percentage of wound closure has increased over both tested cell lines 24 h
after treatment. In terms of antioxidant activity, in a nanosecond pulse radiolysis
and stopped-flow spectrophotometric examination, the ethanol and water extracts of
the stem bark of F. racemosa exhibited significant free radical scavenging activity
in the wound healing process. In addition, the ethanol extract demonstrated consid-
erably more antioxidant potential at steady-state than the water extract.
In addition, Veerapur et al. (2009) conducted an antioxidant assay using conven-
tionally accepted techniques. Using a micronucleus assay on V79 cells, they also
examined the radioprotective ability of the same extract and discovered that the
optimal concentration for radioprotection was 20 g/mL. In the same experiment, the
cytokines did not inhibit the proliferative index. Based on the preceding observa-
tion, the ethanol extract of F. racemosa exhibited substantial antioxidant and radio-
protective traits.

5 Fig Latex and Mechanism of Tissue Repair

Investigation of latex applied externally to wounds reduces inflammation, discom-

fort, and edema and aids healing (Joseph & Raj, 2010). It was noted that several
enzymes, such as trypsin, chymotrypsin, and collagenase, are currently utilized to
heal wounds; these enzymes remove necrotic masses and fibrin clots. It was stated
36 Wound Healing and Ficus carica (Fig) 805

that healing is accelerated when enzymes or proteases are used to treat wounds
(Madhumathi, 2017).
Since antiquity, proteolytic enzymes have been utilized to aid tissue repair. Since
the 1960s, the oral proteolytic enzyme preparation trypsin: chymotrypsin has been
used in clinical settings. It provides superior resolution of inflammatory symptoms
and promotes faster healing of acute tissue injury compared to many currently avail-
able enzyme formulations (Chandanwale et al., 2017). In Ayurveda and Sri Lanka’s
indigenous system of medicine, many portions of Ficus racemosa L., particularly its
latex, have been employed as a wound-healing remedy. Using animal models, this
plant’s wound-healing capacity has been examined (Bopage et al., 2018). Three
sequential but temporally overlapping steps comprise the wound repair process:
inflammation, cell proliferation, and tissue regeneration. The method of wound
healing is fundamentally a connective tissue response. This process begins with an
acute inflammatory phase, followed by the manufacture of collagen and other extra-
cellular macromolecules, which are subsequently reshaped to form scars.
Wound healing active constituents were studied using both in vivo and human
trials; however, now, wound healing active plant secondary metabolites are studied
using both in vitro and in vivo methodologies based on cell cultures of fibroblasts,
keratinocytes, epithelial cells, and endothelial cells (Phan et al., 2000). The break-
ing strength of the incision wounds was significantly raised in the drug-treated
groups, i.e. (257.502.81) in control was increased up to (497.203.33) with F. race-
mosa aqueous extract, and (394.706.61) with F. racemosa ethanol extract. The
results were equivalent to that of the traditional medicine, povidone-iodine, with a
breaking strength. The difference between the drug-treated and control rats was
statistically significant. The aqueous extract ointment treated groups demonstrated
wound healing comparable to that of the reference treatment, povidone-iodine
lotion, beginning on the fourth day. However, with the aqueous extract ointment-
treated group, the wound closure time was shorter, and the percentage of wound
contraction was much higher [12 days for 100% contraction, which was practically
superior to that of the povidone iodine-treated group (17 days)]. Animals treated
with an ointment containing ethanol extract exhibited considerable wound contrac-
tion beginning on the fourth day and wound closure in 14.17 days.
It was reported that the potential wound healing active constituents are the
extracts of stem bark of F. racemose. Lupeol acetate acts as a prodrug for wound
healing through bioactivity-directed fractionation using scratch wound assay (SWA)
over Baby Hamster Kidney (BHK 21) and Maddison-Darby Canine Kidney
(MDCK) epithelial cells. It is recognized that keratinocytes, which create keratin,
are one of the primary cell types that initiate cell proliferation and migration, result-
ing in wound closure. In addition, MDCK cells are capable of generating keratin.
Consequently, even though the MDCK cell line is a broad model of epithelial cells,
it has been utilized as an in vitro model in wound healing assays.
806 N. A. Tayyib

6 Incision and Excision Medel: Wound Healing Activity

The incision and excision models were utilized to investigate the wound healing
activity. In the incision model, the measured parameter was the injured skin’s break-
ing strength. In the excision model, both extracts’ percentage of wound contraction
and duration of epithelialization were determined. The reference standard medicine
for comparison with other groups was povidone iodine ointment. In this model, a
6 cm incision was made through the entire thickness of the skin on either side of the
spinal column of rats. The wound was subsequently closed with 1 cm apart inter-
rupted sutures. The animals were subsequently divided into seven groups at ran-
dom. Rats in Group I (control) received no treatment, rats in Group II (ointment
control) received a topical application of 50 mg of simple ointment base (BP), rats
in Group III were treated topically with standard drug (povidone-iodine), rats in
Group IV were given AQEFR orally (200 mg/kg body weight), rats in Group V were
treated with topical AQEFR ointment (5%), rats in Group VI were given (5%). The
skin’s breaking strength was tested after 10 days (Kokane et al., 2009; Reddy &
Elanchezhian 2008).
In terms of breaking strength in the incision model and percentage of wound
contraction and duration of epithelialization in the excision model, it was deter-
mined that the aqueous extract of F. racemosa had more vigorous wound healing
activity than other groups in both models. In terms of breaking strength in the inci-
sion model and percentage of wound contraction and duration of epithelialization in
the excision model, it was determined that the aqueous extract of F. racemosa had
more vital wound healing activity than other groups in both models (Murti &
Kumar, 2012). In a model of excision wound healing, the aqueous and ethanol
extracts of the root of F. racemosa increase the percentage of wound closure via
promoting epithelialization. This improved epithelialization could result from
F. racemosa extracts’ effect on collagen production. Higher tensile strength indi-
cates faster wound healing. Therefore, it promotes the wound-healing properties of
F. racemosa. The increased tensile strength of treated incision wounds may be
related to increased collagen concentration and fiber stability. Collagen, a compo-
nent of developing cells, is synthesized by a healing tissue.
The platelet-aggregating activity of the aqueous extract of F. racemosa was
investigated in healthy human volunteers. Compared to the controls, the platelet-
aggregating activity of both extracts ranged between 3% and 51%. The dose-
dependent platelet-aggregating activity of both extracts was the same and
dose-dependent. The cold water extract of the bark had bergenin as its primary
chemical ingredient, while the hot water extract also contained ferulic acid, kaemp-
ferol, and coumarin (Ahmed et al., 2012). Numerous extracellular matrix (collagen)
components are synthesized by fibroblast cells, the most frequent cells found in
connective tissue. This study included laboratory mice, separated into control
groups that were not administered in leaf extract and treatment groups that were
administered in the leaf extract. The experimental animals were incised and not
36 Wound Healing and Ficus carica (Fig) 807

administered tin leaf extracts; the control group was examined on days 3 and 7. As
a treatment group, the other group was incised with tin leaf extract and monitored
on days 3 and 7. The extraction of the tin leaf was performed once every day. The
animal was treated with ketamine HCl and diazepam before incision. All incision
wounds on animals used in experiments were treated with wound plaster (hypafix)
to prevent infection and contamination.

7 Evidence Clinical Study of Wound Healing with Fig

The tissue is removed and placed in a sterile tube containing a 10% formalin buffer
solution. The tissue is then placed in liquid paraffin for two to twenty-four hours.
Using a rotary microtome, 4 m-thick paraffin blocks were sliced to a thickness of
4 m. The tissue is stained with hematoxylin and eosin (HE) to determine the number
of fibroblasts and macrophages. In addition, Masson’s Trichome (MT) staining was
carried out to assess the thickness of collagen. When applied to incision wounds, tin
leaf extract increases the quantity of macrophage and fibroblast cells. It denotes a
quickening of macrophage activation and infiltration into the wounded tissue and a
quickening of the inflammatory phase. In addition to macrophage cells, macro-
phages’ production and decreased pro-inflammatory cytokines increased fibroblast
cell proliferation, accelerating wound healing. The extract contains several anti-
inflammatory chemicals, including flavonoids, terpenoids, and tannins.
This plant has been examined by Bopage et al. (2018) to investigate its wound-
healing ability using animal models. This study aimed to acquire insight into the
wound healing process and discover the putative wound healing active substances
present in F. racemosa L. bark utilizing scratch wound assay (SWA) as the in vitro
test. Stem bark extracts of F. racemosa were tested for cell migration boosting abil-
ity and antibacterial activity utilizing the scratch wound assay (SWA) on Baby
Hamster Kidney (BHK 21) and Madin-Darby Canine Kidney (MDCK) cell lines
and the Kirby Bauer disc diffusion assay on common bacteria and fungi.
Dichloromethane and hexane extract that enhanced cell movement on SWA was
fractionated using column chromatography and preparative thin-layer chromatogra-
phy (TLC) to identify the active components. Dichloromethane and hexane extract
enhanced cell migration in both cell lines, while EtOAc and MeOH extracts had
antibacterial and antifungal action against Staphylococcus and Bacillus. It was con-
cluded that chemical components of Ficus racemosa L stem bark stimulate cell
motility and have antibacterial activity. This dual effect supports F. racemosa’s tra-
ditional usage in wound healing. Studies have also shown that the chlorogenic acid
in figs aids in lowering blood sugar and controlling blood glucose levels in type II
diabetes. In diabetic rats, chlorogenic acid (CGA) exhibited antioxidant and pro-
oxidant properties while speeding wound healing (Bagdas et al., 2015).
808 N. A. Tayyib

8 Conclusion

Fig as a medicinal plant that could be scientifically proven for treating wounds was
highlighted in this work. Additional attention has been placed on wound healing in
its natural state, pharmacological activities, the role of fig fruits in wound care, and
metrics used to gauge the effectiveness of those efforts.

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Index

A Antioxidants, 2, 3, 13, 21, 23, 27, 46, 79, 99,

Active compounds, 396, 449, 695, 698 100, 102, 151, 176, 182, 183, 216, 276,
Adaptations, 41, 124, 140, 142, 153, 248, 293–295, 313, 314, 316, 328, 329, 331,
249, 301–303 339, 340, 358, 362, 363, 370, 383–389,
Ames test, 582 396, 397, 403, 406–409, 411, 418, 420,
Anthocyanins, ix, 2, 13, 19, 21, 23, 46, 61, 63, 431, 449–457, 468, 480, 481, 488–490,
64, 96, 98, 99, 151, 156, 182–184, 284, 492, 518, 519, 523, 529, 530, 540, 544,
315, 316, 345–347, 362, 363, 370, 371, 546–552, 562, 563, 565–568, 570–572,
374–377, 380, 382, 384, 389, 396, 398, 583, 587, 598, 626, 630–633, 636, 643,
418, 421, 430, 432, 448–450, 452, 644, 646, 690–696, 698–700, 708, 712,
454–457, 470, 481–488, 503, 514, 540, 733, 734, 757, 758, 788, 789, 793, 795,
544, 548, 563–565, 584, 585, 602, 645, 803, 804, 807
646, 651, 692–694, 696, 699, 733, Applications, vii, ix, x, 2, 4, 7, 27, 52–59, 62,
788, 789 99, 100, 125, 162, 184, 198, 199,
Anticancer, 350, 370, 388, 389, 409, 411, 420, 201–203, 208, 209, 217, 228, 233, 245,
449–452, 468, 488, 489, 491, 546, 549, 249, 267, 271–273, 275, 323, 329, 331,
550, 562, 563, 568, 588, 598, 626, 332, 418, 419, 457, 488, 518, 523–525,
630–633, 636, 692, 695, 803 527–529, 531–533, 539, 548, 571, 585,
Antidiabetic, 322, 370, 408, 409, 411, 452, 453, 620, 621, 623, 628, 630, 643, 644,
489, 513, 519, 523, 540, 545–547, 549, 648–653, 655, 713, 730, 734–738, 746,
551, 563, 567–568, 636, 645, 692, 695 747, 750, 751, 753, 754, 757, 758, 806
Anti-inflammatory, 13, 358, 359, 363, 383, Artificial drying, 792
388, 409, 411, 449, 450, 452, 453, 468,
480, 481, 488–490, 513, 519, 523, 546,
548, 549, 563, 565, 567, 568, 571, 587, B
598, 613, 626, 631, 632, 634, 643, 644, β-caryophyllene, 96, 295, 349, 350, 498–500,
646, 692, 695, 698, 789, 794, 803, 807 515–517, 545, 599, 607
Antimicrobial, 2, 340, 370, 387, 388, 407, Bioactive compounds, ix, 2, 13, 317, 322, 332,
408, 411, 449, 450, 452, 453, 480, 490, 339–341, 346, 348, 362, 387, 389,
491, 518, 519, 546, 549, 551, 626, 628, 396–398, 431, 456, 479–503, 514, 524,
636, 698, 704, 729, 731–736, 738, 739, 528–529, 546, 562–565, 572, 632, 655,
753, 758, 790, 803 690, 691, 694–696, 699, 700, 703,
Anti-mutagenic, 589, 698 712, 737

© The Editor(s) (if applicable) and The Author(s), under exclusive license to 811
Springer Nature Switzerland AG 2023
M. F. Ramadan (ed.), Fig (Ficus carica): Production, Processing, and
Properties, https://doi.org/10.1007/978-3-031-16493-4
812 Index

Bioactivity, 4, 388, 489, 523 Content values, 344, 346, 347, 349–351
Biochemical processes, 730 Coumarins, 96, 261, 295, 340, 347, 387, 398,
Biological activities, 43, 99, 322, 369, 370, 399, 418, 470, 480, 491, 492, 503, 544,
384, 386–389, 397, 411, 418, 419, 450, 547, 549, 584, 585, 598, 600, 602, 605,
453, 454, 488, 517, 519, 523, 550–551, 626, 646, 694, 695, 699, 806
598–636, 645, 753 COVID-19, 29, 30
Brebas, 14, 41, 47, 64, 78, 80, 87, 89, 93, 102, Crops, 29, 41, 42, 44, 45, 47, 48, 56, 64, 78,
109, 110, 112, 114, 115, 119, 120, 80, 87, 89, 102, 103, 109–112, 114,
122–125, 132, 133, 138, 141, 142, 146, 115, 120, 122, 124, 130–133, 137–139,
153, 154, 158, 176, 194, 200, 202, 203, 141, 142, 149, 153, 158, 162, 176, 177,
205, 206, 208, 215, 216, 218, 219, 242, 184–186, 188, 193, 194, 196–198,
243, 252, 733 200–203, 205–208, 215–217, 219, 226,
Breeding, 14, 44, 46–48, 52, 59, 65, 68, 105, 227, 230, 231, 233, 235, 237, 239,
134, 139, 140, 145, 151, 159, 162, 177, 241–243, 248, 249, 252, 262–264, 267,
178, 184, 187, 188, 259–260, 276, 326 269, 271, 273, 274, 277, 285, 292, 325,
Bud, 61, 65, 82, 83, 105, 109–125, 178–180, 326, 342, 382, 396, 480, 497, 562, 572,
194, 202, 241–243, 251, 252, 265, 587, 628, 644, 689, 691, 693, 698, 703,
267, 268 738, 749, 785, 790
By-products, 330, 332, 341, 389, 496, 502, Cultivars, vii, 7, 14, 15, 17–20, 23, 24, 41, 49,
645, 650, 690, 694, 793 50, 59, 78, 80–93, 95, 96, 98, 101–105,
111, 113–115, 117, 120, 124, 130, 132,
134–152, 154, 155, 157–160, 162, 176,
C 183, 184, 194, 197, 200, 202–206,
Caprifigs, 15, 16, 24, 41, 42, 45, 52–56, 64, 215–245, 248–253, 255, 259, 261, 262,
66, 78, 95, 100, 104, 111, 123, 124, 264–266, 271, 272, 276, 284, 292, 293,
129–137, 139, 141, 154, 156–158, 160, 316, 322, 328, 341–346, 349, 363–366,
176–180, 184–188, 250, 260, 261, 263, 372–375, 377, 384, 386, 387, 389, 396,
264, 322, 339, 347, 350, 606–611, 407, 448, 467, 483–487, 491, 496–499,
614–618, 626, 628, 786 501, 502, 514, 515, 518, 519, 524, 525,
Carotenoids, ix, 2, 12, 64, 66, 199, 284, 564, 565, 600, 615, 618, 633, 689, 696,
313–315, 341, 348, 349, 370, 383, 396, 705, 712, 717, 724, 727, 730, 738, 751,
480, 492–494, 498, 503, 514, 564, 566, 753, 754, 788, 795
584, 692, 696, 698, 735 Cultivation, vii, 7, 12, 15, 26, 41, 43, 44, 46,
Cheese making, 622, 746, 747, 754, 755 52, 55, 57, 66, 78, 130, 137, 139, 141,
Chemical characterization, 411 150, 175, 193–209, 215–217, 225, 226,
Chemical composition, 292, 299, 315, 358, 231, 240, 241, 248, 249, 255, 267, 272,
359, 492, 514–515, 517, 569, 795 273, 275, 284, 342, 479, 480, 503, 528,
Chemo-diversity, 21 533, 689, 749, 786, 787, 791
Chitinase, 65, 292, 296, 301, 302, 621, 622,
625, 626, 750, 752
Chocks resilience, 13 D
Climate changes, 29, 104, 105, 252, 253, 270, Defense, 65, 100, 101, 261, 283–303, 403,
330, 533 418, 543, 547, 563, 598, 599, 620, 626,
Climatic conditions, 111, 120, 122, 130, 141, 645, 746, 750
153, 162, 194–198, 247–255, 277, 589, Detection methods, 647, 650, 655–657
689, 786, 787, 790 Deterioration, 197, 244, 248, 690,
Comet assays, 582, 589–591 724, 726–729
Common figs, 40, 41, 43–45, 47, 48, 50–54, Diabetes, ix, 2, 100, 472, 479–481, 489,
56, 57, 59–62, 64–67, 78, 100, 103, 539–553, 562, 567, 568, 570, 585, 633,
109, 110, 112, 123, 124, 129, 135, 145, 692, 705, 707, 789, 807
176, 215, 216, 220, 243, 260, 469, 479, Differentiation, 40, 42, 49, 53, 54, 56, 57, 61,
501, 514, 539, 691, 749, 786, 787 64, 65, 67, 102, 115, 119, 120, 122,
Conservation, 15, 43, 44, 52, 56, 58, 59, 78, 124, 129, 143, 144, 155, 187, 227, 260,
80, 104–105, 130, 134, 162, 726, 299, 598
730, 733 Disease control, 244–245
Index 813

Diversity, vii, 1, 7, 12–21, 23, 24, 27, 31, 292–303, 369, 389, 467, 469, 499, 500,
40–68, 78–105, 130, 132, 134, 135, 526, 539, 545, 546, 549, 550, 581, 583,
137, 139, 141, 144, 145, 148, 159, 162, 585, 590, 598, 622, 634, 645, 692, 746,
177, 178, 180, 181, 183, 186, 255, 267, 748, 749, 803, 805, 807
284, 301, 302, 326, 370, 398, 598, 599, Ficus carica, vii, ix, x, 1–7, 12, 41, 43, 45, 78,
626, 644, 746, 747, 749, 791 80, 87, 93–97, 101, 103, 109, 129–162,
DNA, 24, 45, 47–51, 56, 59, 60, 66–68, 93, 175–188, 193, 204, 217, 227, 247–255,
94, 177, 218, 259, 403, 450, 542–544, 259, 260, 262, 263, 265, 266, 273, 277,
569, 581–583, 589, 590, 631, 632, 737 283, 284, 286–289, 294–298, 303,
DNA fingerprinting, 48, 218 313–317, 339–352, 357–366, 369–389,
Dried figs, 2, 25–29, 48, 79, 98, 102, 137, 138, 396, 397, 399–402, 404–407, 409, 411,
193–196, 260, 267, 269, 276, 277, 418–457, 467, 469, 479–503, 513, 519,
313–317, 331, 347, 358, 360, 363, 523–527, 539–553, 562, 563, 565, 566,
370–375, 384, 388, 397–399, 403, 407, 569–571, 581, 584–591, 599, 620, 634,
408, 410, 411, 421, 450, 451, 453–455, 635, 643–657, 704, 709, 715, 718,
467, 471, 473, 489, 491, 494–501, 503, 724–739, 745–758, 789, 801–808
513, 518, 563, 564, 568, 584, 586, 588, Fig, vii, ix, x, 1–7, 12–31, 40–68, 78–105,
648, 650–653, 657, 690–695, 697, 109, 110, 112–115, 124, 125, 129–162,
707–708, 718, 729, 736, 788, 175–178, 180, 181, 183, 186–188,
790–793, 801 193–209, 215–232, 240–244, 247–255,
259–277, 284, 292, 296, 299, 300, 303,
315, 316, 322–326, 328, 330–332,
E 339–352, 357–366, 369–389, 396–399,
Economic production, 136 407, 410, 421, 431, 432, 448, 451–454,
Ecophysiology, 249–254 456, 457, 467–474, 479–503, 513–519,
Edible figs, 14, 21, 41, 102, 130–133, 136, 523–533, 539, 540, 544–546, 548, 552,
139–160, 176–182, 184–186, 188, 270, 553, 561–573, 583–590, 600, 620–622,
271, 513, 786, 789 625, 626, 630, 631, 644–646, 649–657,
Edible oils, 358 689–700, 703–718, 724–739, 749, 750,
Elicitors, 290, 300 752–755, 785–795, 801–808
Extraction, vii, 7, 99, 201, 237, 260, 328, 331, Fig extracts, ix, 2, 99, 295, 315, 384, 387–389,
332, 339–344, 347, 360–362, 366, 370, 411, 547, 567, 568, 571, 572, 712, 715,
374, 383, 384, 387, 389, 397, 398, 717, 790
400–402, 404–406, 420–433, 457, Fig latex, ix, 1, 66, 244, 261, 339, 341–343,
515–518, 564, 600–603, 605, 614, 616, 351, 431, 518, 524–528, 531, 600,
617, 643–657, 712, 715–717, 746, 602–611, 613–618, 622–624, 626,
747, 807 630–633, 635, 636, 749, 750, 752, 753,
Extraction methods, 374, 383, 384, 397, 398, 755, 756, 758, 802–805
431–433, 514, 516, 519, 643, Fig market, 277
644, 647–656 Fig products, 28, 44, 332, 480, 735
Fig reproductive biology, 132
Flavonoids, 2, 13, 19, 21, 23, 63, 99, 100, 102,
F 151, 183, 199, 294, 295, 313, 315, 340,
Fatty acid contents, 359, 501, 502 341, 343–345, 363, 370, 371, 374–377,
Fatty acids, ix, 2, 43, 98, 100, 102, 323–328, 383, 384, 387, 389, 396, 398, 399,
349–351, 358–360, 470, 480, 500–503, 403–407, 409, 418–449, 451–457, 470,
514, 518, 525, 543, 545, 552, 564, 599, 480, 481, 488, 503, 514, 523, 546–548,
600, 613–619, 625, 789, 794 552, 553, 563–565, 569, 571, 572, 584,
Ficins/ficain, 62, 66, 79, 288, 297, 302, 472, 585, 587, 588, 591, 598, 600–602, 605,
524, 526–533, 568, 584, 589, 599, 626, 631, 633–635, 643, 646, 650, 654,
621–624, 626, 631, 633–636, 746, 747, 656, 657, 691–696, 699, 700, 734, 788,
749–758, 802–804 789, 803, 807
Ficus, 1, 41, 50–52, 78, 79, 95, 103, 124, 129, Food processing, 25, 487, 530, 628–630, 717,
130, 144, 176, 193, 260, 263, 283–289, 736, 747, 750, 791
814 Index

Fruit latex, 261, 525, 588, 600, 621, 623 H

Fruit ripening, 62, 63, 67, 100, 150, 153, 155, Harvesting period, 84, 198, 200, 202–205,
157, 198–199, 202, 208, 220, 221, 244, 207, 717
249, 346, 383, 503, 524, 525, 625, 705, Harvests, ix, 2, 62, 87, 136, 141, 147, 149,
730, 731, 755, 756, 758, 786 154, 156, 181, 195, 198, 200–204, 206,
Fruits, vii, ix, x, 1, 2, 4, 5, 7, 12–24, 26, 27, 31, 208, 209, 241–244, 251, 253, 267, 274,
40–43, 45–47, 50, 54, 56–58, 61–67, 276, 314, 316, 456, 691, 724, 727, 731,
78–80, 84, 87–90, 96–100, 103, 109, 735, 794
110, 112, 114, 115, 122–125, 129–132, Health, vii, ix, 1, 2, 27, 96, 227, 317, 325, 327,
135–142, 144–162, 175–179, 181–188, 359, 370, 384, 388, 397, 403, 408, 418,
193–195, 197–209, 215–225, 227, 228, 420, 455, 470, 472, 479, 492, 494, 496,
230, 231, 240–245, 247–249, 251–253, 498, 501, 503, 513, 519, 523, 533, 553,
260, 261, 263–277, 284, 295, 299, 302, 562, 563, 565, 568, 572, 580, 643, 690,
313–316, 322–325, 327, 328, 330–332, 692, 695, 698–700, 705–708, 726, 746,
340–346, 348–351, 357, 358, 364, 366, 757, 788–790
369–372, 374, 377–379, 381–383, Health benefits, 194, 357, 363, 396, 397,
385–389, 396–402, 404–411, 418, 403–410, 420, 454, 500, 501, 562,
420–445, 448–454, 456, 457, 467–474, 643–645, 654, 657, 690, 692, 695, 698,
479–481, 486–492, 494–497, 499–503, 699, 718
513–517, 519, 523–526, 530, 539–541, Health-promoting attributes, 13, 322, 563
544–551, 562–566, 568–573, 583–591, Health-promoting effect, 43, 363,
599–611, 614–618, 621, 622, 625, 632, 479, 561–573
633, 635, 636, 643–646, 648, 655–657, Health-promoting factors, 13, 322, 561, 790
689–696, 698–700, 703–705, 707–708, High-throughput sequencing (HTS), 59, 67
712, 714–718, 724–738, 746–750, 755, Hydroponics, 201, 233, 235–239
756, 785–795, 801–804, 808
Functional enzyme, 523–533
Functional foods, vii, x, 7, 329, 332, 396, 397, I
562, 630, 692, 698–700, 757, 758, 792 Infections, ix, 2, 264, 271, 276, 291, 292, 297,
Functional traits, 298 301, 302, 471, 532, 544, 553, 626, 628,
630, 636, 727, 789, 790, 802–804, 807
Ingredients, ix, 2, 193, 329, 332, 397, 431,
G 452, 467, 469–471, 580, 697, 706–708,
Genes, 14, 40, 44, 48–50, 56, 59–66, 93, 94, 734, 757, 758, 792–794, 806
100, 101, 123, 124, 140, 150, 155, 157, In vitro study, 524, 802
159, 177, 195, 244, 259, 290, 291, 300,
327, 528, 533, 542, 566, 567, 581, 582,
590, 628 L
Genetic characterization, 43, 44, 79, 80, Latex, vii, ix, 2, 7, 43, 65, 79, 261, 262, 267,
130, 134 287–289, 293, 295–299, 301–303, 322,
Genetic variability, 57, 59, 94, 95, 101, 330, 332, 339, 341–344, 347, 350, 359,
102, 159 374, 387, 388, 409, 421, 426, 431, 432,
Genotoxic, 580–591 448, 457, 472, 500, 514, 516, 518,
Genotypes, 14, 16, 18–24, 43–46, 48, 49, 51, 524–526, 531, 539, 544–546, 553, 563,
52, 55–57, 67, 94, 101, 102, 130, 132, 568, 583–585, 587–589, 598–606,
134, 135, 140, 141, 143, 144, 147, 148, 612–615, 617, 620–636, 746–750,
150–152, 157–162, 177, 178, 180, 181, 752–754, 756–758, 789, 802–805
183, 186, 188, 218, 253, 254, 263, 264, Leaves, vii, ix, 1, 2, 7, 14, 43, 45–47, 50, 61,
276, 298, 299, 341, 345, 373, 375, 65, 79–82, 84, 96, 98–100, 109, 110,
467, 485 113, 114, 125, 131, 140–145,
Glycation, 542, 547–549 158–161, 177–181, 186, 188, 195,
Glycemic index (GI), 705–706 200–202, 205, 216, 218, 227, 228,
Greenhouses, 201–203, 205, 206, 209, 225, 230, 231, 240–242, 244, 245,
226, 229, 231, 235, 377 249–254, 261, 266–271, 284–288,
Index 815

292, 293, 295–299, 301–303, 322, O

339–352, 358, 359, 364, 371–375, Organic acids, ix, 2, 13, 27, 46, 96, 98, 199,
383, 387–389, 408, 409, 421, 424, 238, 284, 313, 362, 369, 398, 469, 480,
426, 427, 429, 431–433, 441–454, 496–498, 503, 514, 523–525, 545, 547,
457, 468–472, 488–491, 496, 497, 599, 633, 692, 693, 695, 698, 788, 792
499–502, 514–517, 519, 523, 524, Oxidative stress, 100, 403, 406, 409, 418, 420,
526, 528, 540, 544, 546–553, 562, 540, 543, 544, 547–549, 552, 567–570,
566–572, 583–588, 590, 591, 631, 635, 636, 644, 789
599–611, 614, 636, 646, 651, 690,
695, 698, 699, 704, 705, 714, 738,
746–749, 789, 793, 802–804, 806, 807 P
Lipochemical attributes, 322, 331, 332 Packaging, 25, 146, 519, 690, 698, 729,
732–733, 736, 739
Pathogens, 65, 101, 150, 195, 199, 261, 284,
M 288–292, 296, 297, 301, 303, 418, 419,
Main crops, 14, 41, 61, 64, 78, 80, 87, 93, 109, 525, 621, 622, 625, 630, 727, 734, 738,
110, 112, 114, 118–120, 122–124, 132, 739, 750, 803
133, 137, 141, 145, 152, 158, 176, 182, Peels, ix, 2, 3, 13, 19, 26, 58, 61, 63, 96, 98,
200, 202, 203, 205–208, 215, 216, 218, 136, 151, 157, 161, 181, 183, 186,
219, 222, 242, 243, 689 284, 315, 330, 332, 342, 344–348,
Mangiferin, 613, 614, 636 350, 351, 371, 372, 385, 386, 389,
Medicinal plants, x, 27, 398, 545, 562, 565, 398, 407, 421, 423, 424, 429,
567, 568, 571, 808 431–433, 438–445, 448–454, 457,
Microbiological spoilage, 724, 726 472, 480, 482, 484–492, 496–499,
Micronucleus assay, 804 501, 502, 514–517, 563–566, 584,
Milk coagulation, 530, 533, 745–758 585, 588, 648, 651–653, 691,
Minerals, ix, 2, 12, 125, 151, 193, 201, 228, 693–695, 700, 724, 734, 735, 788, 793
233–235, 240, 267, 285, 299, 302, 314, Pharmacological evaluation, 411, 546,
358, 369, 388, 396, 456, 467–474, 513, 572, 591
539, 564, 580, 584, 630, 645, 690, 692, Pharmacology, vii, x, 5, 561, 581
695, 698–700, 703, 704, 707, 712, 729, Phenolic acids, 2, 98, 102, 294, 295, 313, 315,
758, 788, 789, 792, 793 316, 340–343, 348, 370, 371, 377, 378,
Modified atmosphere packaging (MAP), 197, 389, 398, 399, 418, 468, 480, 490, 491,
274, 275, 456, 518, 732, 733, 738 501, 503, 514, 525, 544, 548, 563, 564,
Molecular markers, 24, 44, 45, 47, 48, 65, 78, 571, 584, 598, 602, 643, 646, 650, 655,
94, 95, 130, 218 657, 694
Morphological characterization, 79, 80, 134, Phenolic compounds, 23, 96, 98, 99, 102, 156,
140, 145, 146, 157, 159, 162, 186, 187 295, 313, 315, 329, 340–342, 362,
369–371, 374, 375, 377, 382–384,
387–389, 396–411, 418, 432, 433, 455,
N 469, 491, 514, 548, 584, 588, 600,
Natural sun-drying, 792 603–612, 631, 643, 645–648, 651, 653,
Natural sweeteners, 706, 712, 713, 718, 793 655–657, 690, 691, 693–696, 698, 699,
Novel food, x 703, 704, 792
Nutraceutical value, 322, 457, 734 Phenolics, ix, 2, 46, 96, 98–100, 102, 182,
Nutrients, ix, 2, 110, 120, 142, 206, 227, 228, 183, 235, 276, 284, 285, 292–296,
231–238, 266, 267, 290, 297, 314–316, 299, 313–316, 328, 329, 339,
396, 420, 456, 468, 471, 473, 496, 501, 341–343, 358, 362, 369–389,
503, 514, 523, 529, 564, 565, 621, 632, 396–398, 400–402, 407, 418, 429,
645, 690, 691, 697, 698, 703, 707, 712, 430, 432, 433, 470, 490, 492, 514,
729, 734, 736, 786, 793 518, 523, 547–549, 588, 600–602,
Nutrition, vii, x, 7, 150, 193, 201, 231, 273, 623, 626, 633, 636, 643, 645–647,
327, 332, 370, 492, 513, 529, 691, 698, 654, 691–694, 696, 698, 700, 703,
705–707, 758 739, 793
816 Index

Phytochemicals, ix, x, 2, 43, 46, 78, 79, 96–100, Properties, vii, x, 1–7, 26, 43, 46, 50, 54, 80,
151, 183, 184, 188, 284, 314–317, 331, 87, 96, 99, 130, 150, 156, 175–178,
332, 342, 369–371, 377, 388, 389, 396, 183, 187, 188, 218, 232, 267, 272, 275,
398, 400, 418, 480, 514, 539–553, 564, 284, 288, 293, 295, 296, 322, 328, 329,
565, 572, 573, 580, 581, 583–586, 589, 331, 332, 339–352, 358, 359, 363, 366,
591, 598, 630, 631, 636, 643, 644, 647, 369–389, 396, 409, 419, 431, 433, 449,
648, 654–657, 691, 692, 695, 696, 789 450, 452, 453, 468, 480, 481, 490, 496,
Phytosterols, 96, 98, 350, 351, 362, 398, 470, 499, 501, 503, 513, 524, 525, 528–532,
480, 500, 503, 514, 562, 564, 694 545, 546, 548, 549, 562, 563, 567, 568,
Phytotherapy, 586 580, 582, 583, 587–589, 598, 620, 623,
Plant extracts, 27, 374, 375, 549, 568 625, 626, 633, 634, 644–646, 649, 690,
Plant part, vii, 7, 288, 293, 296, 299, 301, 303, 695–699, 706–708, 712, 715–717, 725,
342, 344, 346, 448, 450, 453, 457, 524, 735, 736, 739, 746–749, 751–753,
528, 533, 549, 572, 584, 587–589, 598, 755–758, 786, 788–790, 792, 794,
613, 680, 704, 712, 714 802–804, 806, 807
Plant proteases, 530, 531, 621, 746–748, Proteases, 66, 261, 262, 288, 292, 296–298,
754, 755 302, 524–526, 528, 529, 531–533, 599,
Polymerase chain reaction (PCR), 48, 49, 621–624, 630, 631, 634, 636, 745–756,
58, 61, 94 758, 802–804
Polyphenols, 13, 21, 23, 26, 45, 59, 102, 151, Proteomics, 43, 78, 100–101
176, 183, 296, 302, 363, 370, 371, 384, Pruning intensity, 201–202, 204, 270
387, 397, 398, 467, 470, 480, 481, 503, Pulp, 2, 16, 18, 79, 89, 90, 96, 98, 145,
546, 547, 563, 565, 566, 587, 589, 591, 150–153, 155–158, 179, 182, 185, 186,
633, 634, 644–657, 692, 698–700, 707, 188, 195, 219–224, 243, 264, 315,
708, 734, 788, 789, 792 341–348, 350, 359, 360, 363, 371–373,
Polyunsaturated fatty acids, 98, 100, 325, 327, 375, 378–382, 385–387, 407, 421, 429,
350, 360, 501, 502 431, 438–445, 448, 450–454, 482–492,
Post-harvest valorization, 330 495–499, 501, 514–517, 563–566, 584,
Powder, 193, 260, 340, 344, 346, 691, 693, 588, 648, 651–654, 691, 693–696, 735,
704, 707, 712, 713, 717–718, 791, 793 787, 788
Primordia, 115, 117–120, 123
Processing, vii, ix, x, 1–7, 13, 25, 27, 30, 31,
41, 43, 97, 100, 138, 193, 217, 260, Q
276, 277, 330–332, 341, 359–361, Quality, ix, 2, 13, 21, 23, 25–27, 29, 44, 46,
370, 431, 456, 457, 480, 487, 488, 47, 50, 62, 80, 87, 100, 102, 103, 114,
494, 503, 514, 518, 530, 531, 542, 130, 136, 146, 149–151, 153–155, 160,
569, 572, 583, 645, 655, 690–692, 161, 177, 178, 181, 183, 184, 194, 197,
694, 696, 697, 700, 728, 736, 737, 199, 201, 205, 208, 218, 219, 221–223,
746, 754, 785–795 226, 227, 229, 238, 248, 249, 253,
Processing methods, 693, 700, 790 263–271, 273–277, 284, 322, 327, 370,
Production, vii, x, 1–4, 7, 12, 14, 25, 27, 28, 384, 420, 456, 487, 496, 514, 525, 529,
30, 44, 47, 49, 57, 62, 63, 66, 78, 80, 530, 565, 566, 568, 645, 654, 690, 691,
87, 97, 119, 125, 133, 136–138, 140, 697, 703, 712, 724–738, 746, 756,
146, 153–155, 158, 161, 175, 177, 181, 788, 792–795
182, 184, 194–197, 200, 202–209,
216–218, 225, 226, 230, 231, 233, 244,
247, 248, 253, 261–264, 266, 269, R
271–274, 277, 297, 330, 357, 358, 369, Resilience, 13, 27, 29–31, 253–255, 330, 620
396, 409, 419, 420, 457, 467, 479, 480,
513, 519, 525, 528, 529, 532, 533,
540–542, 544, 547–549, 553, 562, 567, S
573, 580, 622, 625, 631, 634, 635, 689, Seed oil, 43, 322, 324–326, 328, 357,
705, 712–718, 724, 726, 728, 730, 733, 359–363, 502, 552, 567, 794
737, 746, 747, 750, 752, 753, 755, 756, Shelf-life extension, 13, 274, 275, 487, 690,
785–795, 801, 806, 807 729, 730, 733, 739
Index 817

Sterols, ix, 2, 50, 98, 360, 361, 500, 514, 544, V

546, 584, 585, 598, 600, 603–613, Value chain, 13, 14, 27, 29, 30
750, 804 Varieties, ix, 1, 2, 12, 14, 15, 18, 24, 40, 41,
Storage, 13, 25, 29, 30, 62, 104, 145, 275, 285, 43–67, 78–105, 112, 114, 132,
348, 456, 491, 518, 519, 530, 542, 572, 135–140, 148, 150–155, 162, 195, 196,
573, 625, 692, 696, 697, 708, 713, 715, 203, 205, 215, 217, 227, 232, 236, 239,
717, 724, 726–735, 737, 738, 746, 752, 241–243, 260, 265, 266, 271, 292, 299,
753, 792 302, 313–315, 339, 341–348, 350, 351,
Storage conditions, 25, 27, 487, 514, 697, 357, 362, 363, 372, 374–377, 384, 386,
726, 730 389, 396–398, 407, 421, 431, 448, 449,
Sugar, ix, 2, 13, 19, 21, 23, 26, 42, 62, 88, 183, 452, 454, 457, 469, 471, 480, 482,
184, 188, 199, 216, 220, 242–244, 488–490, 492–494, 497, 499, 501, 503,
267–269, 277, 315, 359, 370, 454–456, 518, 541, 563, 565, 566, 568, 625, 628,
470, 487, 513, 523, 525, 527, 530, 632, 643, 657, 691–694, 700, 704, 706,
540–542, 548, 584, 648, 656, 690–692, 712, 725, 749–752, 754, 755, 786–789,
695, 697–700, 703–713, 715, 716, 718, 794, 803
727, 749, 751, 788, 790, 792–795, Vitamins, ix, 2, 12, 100, 193, 276, 313, 314,
801, 807 329, 351, 352, 358, 359, 362, 369, 384,
Sugar substitutes, 705–708, 715 385, 388, 419, 454, 467–471, 480,
Syrup, ix, 1, 27, 267, 330, 331, 436, 457, 530, 494–496, 501, 503, 513, 539, 547, 548,
704, 706, 708, 712–718 553, 564, 566, 584, 630, 645, 692,
695–698, 703, 704, 707, 712, 729, 733,
758, 788, 789, 792, 801
T Volatile compounds, 96, 199, 261, 284, 329,
Tocols, 2 349, 350, 363–366, 398, 498–500, 514,
Transcriptomes, 50, 59, 60, 62–65, 67, 100, 515, 517–519, 523, 525, 788
101, 303, 599, 754
Transcriptomic, 43, 59–67, 78, 100–101
Tree densities, 196, 205, 207 W
Triterpenoids, 96, 98, 294, 295, 398, 480, 503, Wounds, 297, 300, 531, 621, 623,
548, 585, 599, 714 632, 801–808

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Fig (Ficus Carica) : Production, Processing, and Properties: Mohamed Fawzy Ramadan

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Fig (Ficus Carica) : Production, Processing, and Properties: Mohamed Fawzy Ramadan

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Mohamed Fawzy Ramadan

Fig (Ficus carica):

Fig (Ficus carica):

ISBN 978-3-031-16492-7 ISBN 978-3-031-16493-4 (eBook)

Makkah, Saudi Arabia Mohamed Fawzy Ramadan

1 Introduction to Fig (Ficus carica): Production, Processing,

Part I Fig (Ficus carica): Cultivation, Species, and Cultivars

10 Physiological Behaviour of Fig Tree (Ficus carica L.)

Part II Fig (Ficus carica): Chemistry, Functionality

Part III Fig (Ficus carica): Technology, Processing, and Applications

Mohamed Fawzy Ramadan is a Food Chemistry

Mohamed Fawzy Ramadan

1 Fig (Ficus carica): Production, Processing, and Properties

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 1

hydroethanolic extract. In addition, Ficus carica peel extract displayed promising

2 Fig (Ficus carica) Market

3 Fig (Ficus carica) in the Scientific Literature

Note, 12 Short Survey, 1

Fig. 1.3 Distribution by types of the document on Ficus carica. (www.scopus.com)

Muldisciplinary, 28, 1% Neuroscience, 2, 0%

Earth and Planetary Sciences,

Immunology and Microbiology,

Materials Science, 28, 1%

Environmental Science, 124, 5%

Biochemistry, Genecs and

Fig. 1.4 Distribution by subject area of documents on Ficus carica. (www.scopus.com)

Fig. 1.5 Distribution by country of documents on Ficus carica. (www.scopus.com)

4 Aims and Features of This Book

Lahcen Hssaini, Rachid Razouk, Aziz Fadlaoui, and Karim Houmanat

ABTS Ethylbenzothiazoline-6-sulfonic acid

L. Hssaini (*) · R. Razouk · A. Fadlaoui · K. Houmanat

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 11

Morocco hosts large biodiversity and considerable within-species genetic diversity

(lycopene as the most abundant, zeaxanthin, α-carotene, lutein, cryptoxanthin, and

2 Genetic Diversity Patterns

2.1 Overview of Available Fig Germplasm

Table 2.1 Denomination meaning of some fig cultivars in Morocco

Hssaini et al. (2019b)

environmental effect, this confusion in the cultivar denomination is linked to the

2.2 Fruit Morphometric Diversity

Table 2.2 (continued)

However, according to the International Commission on Illumination (CIE), the

Moreover, both browning types speed up as the temperature increases (Bet,

4 Value Chain and Its Resilience

oligopolistic nature, characterized by numerous small and medium-sized producers,

investment in new agricultural technologies. These weaknesses can broadly weaken

El Khaloui, M. (2010). Valorisation de la figue au Maroc. Transfert de Technologie en Agriculture,

Dunja Bandelj, Alenka Baruca Arbeiter, and Matjaž Hladnik

D. Bandelj (*) · A. Baruca Arbeiter · M. Hladnik

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 39

n Number of amplified alleles

1 Origin, Taxonomy and Biological Properties

2 Fig Genetic Resources: The Importance of Research

3 Tools for Fig Genetic Resources Diversity Studies

The traditional approach of fig germplasm characterization is based on the descrip-

The development of DNA markers has further enriched the characterization

3.2.1 Development of Fig Microsatellites

Microsatellites, or simple sequence repeats (SSRs), have an important place in plant

sequences in microsatellite flanking regions and evolution distance of species may

3.3 Application of Microsatellites in Fig Diversity Studies

Table 3.1 (continued)

agroecosystem. This spontaneous hybridization represents an important driving

4 Gene, Genomic and Transcriptomic Studies of Figs

Technologies for high-throughput sequencing (HTS) have enabled the generation of

The common fig is a diploid species with 2n = 26 chromosomes (Flaishman et al.,

4.1.2 New Research Approaches and Findings Based

genetic map. RAN1 is involved in the transduction of ethylene, and a suggested

4.2 Transcriptomic and Gene-Based Studies

4.2.1 Mechanisms Related to Fruit Development and Ripening

To elucidate biological mechanisms related to plant and fruit development, several

using Agrobacterium-mediated transformation of a construct carrying the CRISPR/

Transcriptome studies related to the flavonoid biosynthetic pathways were con-

4.2.3 Physiological Differences Between Common Fig Types

also differentially expressed. A study of FcFT1, a FLOWERING LOCUS T, was

4.2.4 Response to Abiotic Factors

4.2.5 Response to Biotic Factors

Makkah, Saudi Arabia Mohamed Fawzy Ramadan

1 Introduction to Fig (Ficus carica): Production, Processing,

10 Physiological Behaviour of Fig Tree (Ficus carica L.)

Part II Fig (Ficus carica): Chemistry, Functionality

1 Fig (Ficus carica): Production, Processing, and Properties

2 Fig (Ficus carica) Market

3 Fig (Ficus carica) in the Scientific Literature

4 Aims and Features of This Book

2 Genetic Diversity Patterns

2.1 Overview of Available Fig Germplasm

2.2 Fruit Morphometric Diversity

4 Value Chain and Its Resilience

1 Origin, Taxonomy and Biological Properties

2 Fig Genetic Resources: The Importance of Research

3 Tools for Fig Genetic Resources Diversity Studies

3.2.1 Development of Fig Microsatellites

3.3 Application of Microsatellites in Fig Diversity Studies

4 Gene, Genomic and Transcriptomic Studies of Figs

4.1.2 New Research Approaches and Findings Based

4.2 Transcriptomic and Gene-Based Studies

4.2.1 Mechanisms Related to Fruit Development and Ripening

4.2.3 Physiological Differences Between Common Fig Types

4.2.4 Response to Abiotic Factors

4.2.5 Response to Biotic Factors

Blastophaga psenes L. Evolution, 41(4), 693–704. https://doi.org/10.1111/j.1558-5646.1987.

2 Genetic Variability in Fig Germplasm

2.4 Molecular and Biochemical Variability Assessment

2.4.1 Structure and Composition of Fig Genome

2.5 Biochemical Traits and Phytochemical Contents Divers

2.6 Proteomic and Transcriptomic Analysis

3 Multivariate Statistical Analysis of Qualitative

4 Variation in Graft Compatibility of Wild Fig Species

5 Trends in Fig Germplasm Conservation

1 Fig Buds Description

2 Development and Evolution of Inflorescence

3 Influence of Cultural Practices and Plant Growth

2 Botany and Biology in Different Types of Figs

3 Germplasm Diversity and Conservation

4 Wild Figs, Cultivars, Local Clones, and Hybrids

4.2 Cultivars and Local Clones

4.3 Hybrids and Transgenic Varieties

5.3.1 Environmental Condition’s Effects on Fruit Characteristics

5.3.2 Caprification Effects on Fruit Morphological Characteristics

6 Descriptor’s Optimization Tools

6.1 Principal Component Analysis (PCA)

2 Plant and leaf Characteristics