Sage The Genus Salvia - 1st Edition

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This edition published in the Taylor & Francis e-Library, 2005.

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Copyright © 2000 OPA (Overseas Publishers Association) N.V. Published by license

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Copyright © 2000 OPA (Overseas Publishers Association) N.V. Published by license under the
Harwood Academic Publishers imprint, part of the Gordon and Breach Publishing Group.
 CONTENTS

Preface to the Series vii

Preface ix
Contributors xv
Acknowledgements xix

I. Introduction
1. The Folklore and Cosmetic Use of Various Salvia Species 1
Anthony C.Dweck

II. Botany
2. The Sage Plants in Greece: Distribution and Infraspecific Variation 27
Regina Karousou, Effie Hanlidou and Stella Kokkini
3. Salvia in Southern Africa 47
Anna K.Jäger and Johannes van Staden

III. Chemical Constituents

4. Terpenoids in the Genus Salvia 55
Ayhan Ulubelen
5. Essential Oils of Salvia spp: Examples of Intraspecific and
Seasonal Variation 69
Amalia L.Giannouli and Spiridon E.Kintzios
6. Salvianolic Acids and Related Compounds 81
Lian-Niang Li

IV. Cultivation and Breeding

7. The Cultivation of Sage 93
Andreas J.Karamanos
8. Genetic Improvement of Cultivated Species of the Genus Salvia 109
Jenõ Bernáth and Éva Németh
9. The Ecophysiology of Salvia: Disorders and Adaptation 125
Emmanouil Panagiotopoulos, Chrisostomos Kapetanos, Maria Skapeti,
Constantinos Cholevas, John Drossopoulos, Michael Loukas and
Spiridon E.Kintzios
10. Legal Protection of Salvia Varieties 135
Bernard Le Buanec
V. Pharmacology
11. The Biological/Pharmacological Activity of the Salvia Genus 143
Dea Baricevic and Tomaz Bartol
12. Antioxidants from Salvia officinalis 185
Stanley G.Deans and Elisabeth J.M.Simpson

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vi CONTENTS

13. The Antihypertensive Properties of Dan-Shen, the Root of

Salvia miltiorrhiza 193
Takako Yokozawa
14. Why Sage May be a Wise Remedy: Effects of Salvia on the Nervous
System 207
Nicolette Perry, Melanie-Jayne Howes, Peter Houghton and Elaine Perry

VI. Biotechnology
15. The Biotechnology of Labiatae 225
Olga Makri
16. In vitro Rosmarinic Acid Production 233
Isabelle Hippolyte
17. Salvia spp.: Tissue Culture, Somatic Embryogenesis, Micropropagation
and Biotransformation 243
Spiridon E.Kintzios
18. Production of Secondary Metabolites using Liquid Culture of Salvia
Plants: Up-to-Date Reports and Scale-up Potential 251
Emmanouil Panagiotopoulos, Maria Skapeti and Chrisostomos
Kapetanos

VII. Commercial Aspects

19. Production of Salvia Oil in Mediterranean Countries 263
K.Husnu Can Baser

VIII. General Issues

20 Scientometric Analysis of Science and Technology Bibliographic
Information Sources with Regard to Genus Salvia 269
Tomaz Bartol and Dea Baricevic

Appendix
Short Notes on the Olfactory Properties of the Essential Oil of Some Sage
Species 289

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Harwood Academic Publishers imprint, part of the Gordon and Breach Publishing Group.
 PREFACE TO THE SERIES

There is increasing interest in industry, academia and the health sciences in medicinal
and aromatic plants. In passing from plant production to the eventual product used
by the public, many sciences are involved. This series brings together informa tion
which is currently scattered through an ever increasing number of journals. Each
volume gives an in-depth look at one plant genus, about which an area specialist has
assembled information ranging from the production of the plant to market trends
and quality control.
Many industries are involved such as forestry, agriculture, chemical, food,
flavour, beverage, pharmaceutical, cosmetic and fragrance. The plant raw materials
are roots, rhizomes, bulbs, leaves, stems, barks, wood, flowers, fruits and seeds.
These yield gums, resins, essential (volatile) oils, fixed oils, waxes, juices, extracts
and spices for medicinal and aromatic purposes. All these commodities are traded
worldwide. A dealer’s market report for an item may say “Drought in the country of
origin has forced up prices”.
Natural products do not mean safe products and account of this has to be taken
by the above industries, which are subject to regulation. For example, a number of
plants which are approved for use in medicine must not be used in cosmetic
products.
The assessment of safe to use starts with the harvested plant material which has to
comply with an official monograph. This may require absence of, or prescribed
limits of, radioactive material, heavy metals, aflatoxins, pesticide residue, as well as
the required level of active principle. This analytical control is costly and tends to
exclude small batches of plant material. Large scale contracted mechanised
cultivation with designated seed or plantlets is now preferable.
Today, plant selection is not only for the yield of active principle, but for the
plant’s ability to overcome disease, climatic stress and the hazards caused by
mankind. Such methods as in vitro fertilisation, meristem cultures and somatic
embryogenesis are used. The transfer of sections of DNA is giving rise to controversy
in the case of some end-uses of the plant material.
Some suppliers of plant raw material are now able to certify that they are
supplying organically-farmed medicinal plants, herbs and spices. The Economic
Union directive (CVO/EU No 2092/91) details the specifications for the obligatory
quality controls to be carried out at all stages of production and processing of
organic products.
Fascinating plant folklore and ethnopharmacology leads to medicinal potential.
Examples are the muscle relaxants based on the arrow poison, curare, from species
of Chondrodendron, and the antimalarials derived from species of Cinchona and
Artemisia. The methods of detection of pharmacological activity have become
increasingly reliable and specific, frequently involving enzymes in bioassays and

vii
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viii PREFACE TO THE SERIES

avoiding the use of laboratory animals. By using bioassay linked fractionation of

crude plant juices or extracts, compounds can be specifically targeted which, for
example, inhibit blood platelet aggregation, or have antitumour, or antiviral, or any
other required activity. With the assistance of robotic devices, all the members of a
genus may be readily screened. However, the plant material must be fully
authenticated by a specialist.
The medicinal traditions of ancient civilisations such as those of China and India
have a large armamentarium of plants in their pharmacopoeias which are used
throughout South East Asia. A similar situation exists in Africa and South America.
Thus, a very high percentage of the world’s population relies on medicinal and
aromatic plants for their medicine. Western medicine is also responding. Already in
Germany all medical practitioners have to pass an examination in phytotherapy
before being allowed to practise. It is noticeable that throughout Europe and the
USA, medical, pharmacy and health related schools are increasingly offering training
in phytotherapy.
Multinational pharmaceutical companies have become less enamoured of the
single compound magic bullet cure. The high costs of such ventures and the endless
competition from too many compounds from rival companies often discourage the
attempt. Independent phytomedicine companies have been very strong in Germany.
However, by the end of 1995, eleven (almost all) had been acquired by the multi
national pharmaceutical firms, acknowledging the lay public’s growing demand for
phytomedicines in the Western World.
The business of dietary supplements in the Western World has expanded from the
Health Store to the pharmacy. Alternative medicine includes plant based products.
Appropriate measures to ensure the quality, safety and efficacy of these either
already exist or are being answered by greater legislative control by such bodies as
the Food and Drug Administration of the USA and the recently created European
Agency for the Evaluation of Medicinal Products, based in London.
In the USA, the Dietary Supplement and Health Education Act of 1994 recognised
the class of phytotherapeutic agents derived from medicinal and aromatic plants.
Furthermore, under public pressure, the US Congress set up an Office of Alternative
Medicine and this office in 1994 assisted the filing of several Investigational New
Drug (IND) applications, required for clinical trials of some Chinese herbal prepa
rations. The significance of these applications was that each Chinese preparation
involved several plants and yet was handled as a single IND. A demonstration of the
contribution to efficacy, of each ingredient of each plant, was not required. This was
a major step forward towards more sensible regulations in regard to
phytomedicines.
My thanks are due to the staff of Harwood Academic Publishers who have made
this series possible and especially to the volume editors and their chapter
contributors for the authoritative information.

Roland Hardman

Copyright © 2000 OPA (Overseas Publishers Association) N.V. Published by license under the
Harwood Academic Publishers imprint, part of the Gordon and Breach Publishing Group.
 PREFACE

Salvia is a fascinating plant genus. One of the widest-spread members of the Labiatae
family, it features prominently in the pharmacopoeias of many countries throughout
the world. From the Far East, through Europe and across to the New World several
of the almost 1000 Salvia species have been used in many ways, e.g. essential oils
used in perfumery, the flowers used as rouge, the leaves used for varicose veins, the
seed oil as an emollient, the roots as a tranquiliser. The range of traditional
applications of the herb in domestic medicine seems to be endless: it has been used as
a medication against perspiration and fever; as a carminative; a spasmolytic; an
antiseptic/bactericidal; an astringent; as a gargle or mouthwash against the
inflammation of the mouth, tongue and throat; a wound-healing agent; in skin and
hair care; and against rheumatism and sexual debility in treating mental and nervous
conditions as well as an insecticidal.
This book begins with the presentation of the (approximately 400) most known
Salvia species (Chapter One, A.C.Dweck), their pharmacopoeial status, their history
and distribution, traditional uses as a food source and in domestic medicine, as well
as general information on the chemical composition of prominent Salvia species,
such as S. officinalis, S. bowleyana, S. coccinea, S. columbariae, S. digitaloides, S.
divinorum, S. hispanica, S. horminum, S. lavandulaefolia, S. miltiorrbiza, S. plebeia,
S. pomifera, S. repens, S. rugosa, S. runcinata, S. sisymbrifolia, S. sclarea, S. erotina,
S. verbenaca and S. yunnanensis. Analytical dosing instructions are given for each
area of application.
The botany and the distribution—both global and regional—of the genus is
presented along with taxonomical, chemotaxonomical, genetical and phylogenical
aspects. In Chapter Two (R.Karousou et al.) detailed information is provided on sage
species growing in Greece and the ad hoc main centre of origin and native
distribution, the Mediterranean region. Emphasis is given on the three main species
endemic in the region, namely S. officinalis (‘Dalmatian or Garden Sage’), S.
fruticosa (‘Greek Sage’) and S. pomifera (‘Cretan Sage’), providing detailed
information on their geographic distribution, morphology and essential oil
composition. These species are remarkably variable and there is a vivid presentation
of the climatically (temperature- and precipitation-) related high variation of the leaf
mor phology and the qualitative and quantitative essential oil content (due, for
example, to xerophytic adaptation). On the opposite side of the globe, Southern
Africa, where traditional medicine plays a very important role in health care, is home
to 30 species of the genus Salvia. In Chapter Three (A.K.Jäger and J.van Staden) we
learn about the botany, the distribution in different climatic regions, the traditional
usages and the chemistry of representative Salvia species, like S. africana, S.
chamelaeagmea and S. stenophylla, the latter species being one of the few known
sources of epi-a-bisabolol, a potent anti-inflammatory agent.

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x PREFACE

An unusually large number of useful secondary metabolites, belonging to various

chemical groups, have been isolated from Salvia species. These include various
phenolic acids (such as caffeic, chlorogenic, ellagic, ferulic and gallic acid), tannins
and volatile substances. Chapters 4–6 provide detailed information on the
extraction, isolation and characterization of those components to which the
biological properties of sage can be attributed: terpenoid compounds, essential oils
in general and phenolic derivatives like salvianolic acids (including rosmarinic acid
and lithospermic acid). In Chapter Four (A.Ulubelen), the presence of terpenoids
(except of monoterpenoids) in Salvia species is thoroughly discussed. The chemical
structure and botanical distribution and, in certain cases, structure-related bioactive
properties of a total of 111 terpenoids is discussed, including various groups of
diterpenoids (abietane, clerodane, pimarane and labdane-type), triterpenoids,
sesquiterpenoids and sesterterpenoids. Cumulative data on the seasonal and
intraspecific variation of essential oils are also included (Chapter Five, A.L.
Giannouli and S.E.Kintzios), indicating that it may be possible to manipulate
essential oil content in such a way that heavy investment of time and resources in
selection and breeding can be avoided. In Chapter Six (Lian-Niang Li) the chemistry
of the bioactive polyphenolic acids of various Salvia species, in particular S. miltior
rhiza, is presented. These substances are commonly known as rosmarinic acid,
lithospermic acid, salvianolic acids (A-J) and related compounds. Detailed
information is provided on their chemical structure, extraction and isolation
methods, UV-, MS- and NMR-spectra and chemical transformation.
The optimization of tillage, harvest and dry process technologies, as well as the
proper application of fertilizers and pesticides largely improved the quality of the
raw material. Chapter Seven (A.J.Karamanos) constitutes an in-depth review of
virtually every aspect concerning the cultivation technology of sage, such as propa
gation, land preparation, irrigation, fertilizer, growth regulator and herbicide
application, harvest and postharvest treatment. A special reference is made to the
ecophysiology of the genus in respect of its response to abiotic and biotic stress
factors (drought, heavy metals, light, temperature, allelopathy, etc.) and their effect
on biomass production and product yield and quality (e.g. essential oil composition)
(Chapter Nine, E.Panagiotopoulos et al.).
Regarding optimization of the cultivation of the species for appropriate plant
material, initially only indigenous local or introduced populations were used with
moderate efficacy. To provide the basis for economical production, the breeding
work especially on S. sclarea and S. fficinalis became more intensive. Breeding work
on the genus is currently done in some countries (mainly Eastern European ones) to
obtain varieties with improved characteristics, in their agricultural behaviour as well
as in their chemical composition. In Chapter Eight (J.Bernáth and É.Németh) we see
how various selection goals are approached by different strategies and methods
country by country, depending on the local tradition and experiences. The high
morphological and chemical diversities of the species are utilised, even if the plant is
growing wild locally, or the indigenous populations had been introduced from
exterior habitats. In particular there is extensive reference to the utilization of
morphological and chemical diversity of indigenous populations as a background

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 PREFACE xi

for genetic improvement, the improvement of populations by selection, the creation

of new cultivars by hybridisation, the construction of polyploid forms and mutation
breeding. A continuous breeding effort, incorporating both classical and modern,
unconventional methods, led to the creation of new productive varieties (such as
‘Extrakta’, a S. officinalis cultivar which is particularly rich in essential oil). Breeding
new sage varieties requires substantial investment in terms of skill, labour, material
resources and money and may take years. An efficient protection of the intellectual
property should enhance the breeding work on the Salvia genus. The 1991 Act of the
UPOV Convention and the 1994 TRIP’S Agreement provide for the possibility of
protecting all genera and species in many countries. Hence, this gives a new
opportunity to breeders of ornamental, medicinal and aromatic plants and in
particular to Salvia breeders. The current situation of protection with a detailed
explanation of the UPOV plant protection system and its main dispositions are given
in Chapter Ten (B.Le Buanec).
The biological effects of plant extracts and/or essential oils and other important
compounds of various species of the genus Salvia have been acknowledged over the
centuries. Besides S. officinalis, which is additionally referred to as having antibiotic
properties, other species also contain compounds with important pharmaco logical
activities. In Chapter Eleven (D.Baricevic and T.Bartol), the bioactive/
pharmacological properties of the genus are quite extensively reviewed, including
antimicrobial, antiviral, cardiovascular, renal, antioxidative, anti-inflammatory,
tumorigenesis-preventing, antimutagenic, peptic-antiulcer, antispasmodic,
hypoglycemic and hepatoprotective activities, as well as documented toxic effects.
Extension of the use of sage as a food additive or a herbal medicine has been
prevented mostly due to the toxic effects of the ketone terpenoids in the volatile oil,
namely camphor and thujone. A separate section of this chapter is devoted to the
description of the pest-toxic and repellent activities of the genus.
Because of the unknown effect of synthetic antioxidants like butylated
hydroxyanethole [BHA] and butylated hydroxytoluene [BHT] on human cancer risk
the interest in preparing antioxidants from natural sources with minimal processing
has considerably increased in recent years. Sage and related species are an important
source of antioxidants used in the food industry and have wider implications for the
dietary intake of natural antioxidants. Salvia is one of the favourite candidate species
as a source of natural antioxidants in health care products. In Chapter Twelve
(S.G.Deans and E.J.M.Simpson) the chemical structure, isolation and activity of the
major antioxidant compounds of sage, mainly rosmarinic acid, carnosol, carnosic
acid, rosmadial, rosmanol, epirosmanol and methyl carnosate, are presented.
Aspects of essential oil variability and herbal material purity due to geographic
location and drying temperature are investigated, along with a short description of
assays for the determination of antioxidant activity. There is an analytical reference
to studies investigating the relationship between leaf senescence and the oxidative
defense system of sage, as well as the activity of the antioxidants in improving the
responsiveness of the human immune system and reducing the free radical damage.
Dan-Shen extracts, derived from the roots of S. miltiorrhiza, have traditionally
been used to treat haematological abnormalities and cardiovascular diseases in

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xii PREFACE

China. Along with tanshinone IIA sodium sulfonate, magnesium lithospermate B (a

tetramer of caffeic acid) is an important constituent with antihypertensive
properties. In Chapter Thirteen (T.Yokozawa) the structure and activity of this and
related compounds is evaluated in detail, presenting its effects on blood flow and
renal function as a result of the interaction with the secretion of prostaglandin E2 and
kallikrein activation.
Sage is also renowned for its effects on the central nervous system. Various
anxiolytic and sedative, memory-enhancing, antidepressive and hallucinogenic
activities have been ascribed to the genus (Chapter Fourteen, N.Perry et al.), thus
making it a promising ingredient in the future treatment of CNS-related ailments.
Biotechnological techniques have been recently reported to significantly facilitate
plant propagation and production of some important bioactive compounds of the
genus Salvia. A special chapter of the book (Chapter Fifteen, O.Makri) is devoted to
reviewing the biotechnology research that has been done in various other species of
the Labiatae family, such as Mentha, Origanum, Thymus, Lavantula, Ocimum,
Hyssopus and Coleus, whose antioxidant activity is valuable for the food, cosmetic
and pharmaceutical industries.
Rosmarinic acid can be produced by cell suspension cultures of sage. The growth
and production of rosmarinic acid by sage cells is modified by the type of culture
medium used (Chapter Sixteen, I.Hippolyte). Rosmarinic acid production is
increased 10-fold to attain 6.4 gL-1 (or 36% of the dry weight) under optimal
conditions. Investigation of cell growth kinetics showed that a change in the medium
caused shifts in peaks of growth and rosmarinic acid production, and modifications
in cellular metabolism. By changing the composition of the culture medium it is
possible to manipulate rosmarinic acid production to coincide with cell growth or to
begin only when growth had stopped.
There is an increasing interest in the development of efficient protocols for the
tissue culture and micropropagation of certain Salvia species, in order to establish a
relatively fast system for producing disease-free and true-to-type clonal (and
therefore uniform) plants from outstanding genotypes. Progress in somatic
embryogenesis and recent research on the technology of synthetic seeds, along with
other advanced aspects of tissue culture (e.g. protoplast culture and fusion, creation
of autotetraploid lines) could offer a significant involvement of biotechnology in the
propagation and breeding of the genus Salvia.
Chapter Seventeen (S.E.Kintzios) offers a concise presentation of the various
methods developed for the induction of callus, organogenesis and somatic
embryogenesis as well as plant regeneration for micropropagation and breeding
purposes of some Salvia species. Furthermore, the accumulation of secondary
metabolites (in particular rosmarinic and lithospermic acid) in in vitro differentiated
tissues is reviewed.
Biotechnology opens new perspectives for an automated, scaled-up and cost-
efficient production of useful compounds from Salvia spp. Cell suspensions, immob
ilized cell and hairy root cultures have been established from S. officinalis, S.
miltiorrhiza, S. fruticosa and S. sclarea and used for the production of various
secondary metabolites, such as rosmarinic acid, cryptotanshinone, ferruginol

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 PREFACE xiii

(achieving a yield of 29, 101 and 254 g/1, respectively) and (commercially) sclareol.
Scale-up and immobilization techniques for Salvia liquid cell culture are the focus of
Chapter Eighteen (E.Panagiotopoulos et al.), where updated information on the in
vitro secondary metabolism of various important compounds can also be found.
It is hard to describe sage as an industrial crop, since its worldwide production is
less than 25 000 kg per year. There is, however, a steady upward trend in the export
of essential oil from various sage species, for use as products in the aromatherapy
and natural cosmetics market. Commercial sage species include S. officinalis, S.
fruticosa, S. lavandulaefolia, S. verbenaca, S. sclarea and S. tomentosa. In Chapter
Nineteen (K.H.C.Baser) the actual situation of sage oil production and export
(mainly in Mediterranean countries) is presented. In Turkey alone, approximately
500 kg of leaf oil from S. triloba (S. fruticosa) is annually produced and 600 tonnes
of sage leaves worth more than 1.5 million US$ is exported. There is, in addition, an
increasing demand for herbal tea from organically grown sage.
In the age of information technology a researcher is facing an accelerating growth
of all kinds of scientific and technical data. The average yearly growth of Salvia-
related publications is 2.2%. This rate is slight but persistent and shows a con
tinuing presence of interest in the field. In an exhaustive analysis (Chapter Twenty,
T.Bartol and D.Baricevic) most major bibliographic life-sciences databases are
identified and assessed as pertinent sources for information on the genus Salvia.
Using the example of Salvia, the degree of overlap across databases is observed and
the annual trend of publishing for this genus and major journals where Salvia-related
articles have been published is identified. An investigation of these databases and
their differences in relation to keyword or classification representation of the topics
is conducted. Finally, the most appropriate search technique in order to maximize
the recall and optimize the precision is selected and presented.

Spiridon E.Kintzios

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Harwood Academic Publishers imprint, part of the Gordon and Breach Publishing Group.
 Dedicated to Katia

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Harwood Academic Publishers imprint, part of the Gordon and Breach Publishing Group.
 CONTRIBUTORS

Dea Baricevic Constantinos Cholevas

Slovenian National AGRIS Centre Department of Plant Physiology
Biotechnical Faculty Faculty of Agricultural Biotechnology
University of Ljubljana Agricultural University of Athens
Jamnikarjeva 101 Iera Odos 75
1111 Ljubljana 11855 Athens
Slovenia Greece
Tomaz Bartol Stanley G.Deans
Slovenian National AGRIS Centre Aromatic and Medicinal Plants Group
Biotechnical Faculty SAC Auchincruive
University of Ljubljana Ayr KA6 5HW
Jamnikarjeva 101 United Kingdom
1111 Ljubljana
Slovenia John Drossopoulos
Department of Plant Physiology
K.Husnu Can Baser
Faculty of Agricultural Biotechnology
Anadolu University
Agricultural University of Athens
Medicinal and Aromatic Plant and Drug
Iera Odos 75
Research Centre (TBAM)
11855 Athens
Yunus Emre Kampusu
Greece
26 470 Eskisehir
Turkey Anthony C.Dweck
Jeno Bernáth Peter Black Medicare Ltd.
Department of Medicinal Plant White Horse Business Park
Production Aintree Avenue
University of Horticulture and Food Trowbridge
Industry Wiltshire BA14 0XB
Villanyi Str. 29/31 United Kingdom
Budapest
Amalia L.Giannouli
Hungary
Department of Plant Physiology
Bernard Le Buanec Faculty of Agricultural Biotechnology
International Association of Plant Agricultural University of Athens
Breeders (ASSINSEL) Iera Odos 75
Chemin du Reposoir 7 11855 Athens
1260 Nyon Greece
Switzerland

xv
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xvi CONTRIBUTORS

Effie Hanlidou Andreas J.Karamanos

Laboratory of Systematic Botany and Laboratory of Crop Production
Phytogeography Faculty of Crop Science and Production
School of Biology, Faculty of Sciences Agricultural University of Athens
Aristotle University Iera Odos 75
54006 Thessaloniki 11855 Athens
Greece Greece

Isabelle Hippolyte Regina Karousou

7, place Albert 1er Laboratory of Systematic Botany and
34 000 Montpellier Phytogeography
France School of Biology, Faculty of Sciences
Aristotle University
Peter Houghton 54006 Thessaloniki
Pharmacognosy Research Laboratories Greece
Pharmacy Department
King’s College London Spiridon E.Kintzios
Manresa Road Department of Plant Physiology
London SW3 6LX Faculty of Agricultural Biotechnology
United Kingdom Agricultural University of Athens
Iera Odos 75
Melanie-Jayne Howes 11855 Athens
Pharmacognosy Research Laboratories Greece
Pharmacy Department
Stella Kokkini
King’s College London
Laboratory of Systematic Botany and
Manresa Road
Phytogeography
London SW3 6LX
School of Biology, Faculty of Sciences
United Kingdom
Aristotle University
Anna K.Jäger 54006 Thessaloniki
Research Unit for Plant Growth and Greece
Development Lian-Niang Li
Department of Botany Institute of Materia Medica
University of Natal Pietermaritzburg Chinese Academy of Medical Sciences
Private Bag X01 Peking Union Medical College
Scottsville 3209 1 Xian Nong Tan Street
South Africa Beijing 100050
China
Chrisostomos Kapetanos
Department of Plant Physiology Michael Loukas
Faculty of Agricultural Biotechnology Department of Genetics
Agricultural University of Athens Faculty of Agricultural Biotechnology
Iera Odos 75 Agricultural University of Athens
11855 Athens Iera Odos 75
Greece 11855 Athens
Greece

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 CONTRIBUTORS xvii

Olga Makri London SW3 6LX

Department of Plant Physiology United Kingdom
Faculty of Agricultural Biotechnology
Elisabeth J.M.Simpson
Agricultural University of Athens
Aromatic and Medicinal Plants Group
Iera Odos 75
SAC Auchincruive
11855 Athens
Ayr KA6 5HW
Greece
United Kingdom
Éva Németh
Maria Skapeti
Department of Medicinal Plant
Department of Plant Physiology
Production
Faculty of Agricultural Biotechnology
University of Horticulture and Food
Agricultural University of Athens
Industry
Iera Odos 75
Villanyi Str. 29/31
11855 Athens
Budapest
Greece
Hungary
Ayhan Ulubelen
Emmanouil Panagiotopoulos
Faculty of Pharmacy
Department of Plant Physiology
University of Istanbul
Faculty of Agricultural Biotechnology
34 452 Istanbul
Agricultural University of Athens
Turkey
Iera Odos 75
11855 Athens Johannes van Staden
Greece Research Unit for Plant Growth and
Development
Elaine Perry
Department of Botany
Medical Research Council (MRC)
University of Natal Pietermaritzburg
Neurochemical Pathology Unit
Private Bag X01
Newcastle General Hospital
Scottsville 3209
Westgate Road
South Africa
Newcastle-upon-Tyne NE4 6BE
United Kingdom Takako Yokozawa
Research Institute for Wakan-Yaku
Nicolette Perry
Toyama Medical and Pharmaceutical
Pharmacognosy Research Laboratories
University
Pharmacy Department
2630 Sugitani
King’s College London
Toyama 930–0194
Manresa Road
Japan

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