See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/353292865

Bougainvillea

Chapter · July 2021

DOI: 10.1007/978-981-15-1554-5_2-1

CITATION READS
1 3,091

3 authors, including:

Subodh Kumar Datta Datta T. Janakiram

Retired 58 PUBLICATIONS 326 CITATIONS
156 PUBLICATIONS 2,545 CITATIONS
SEE PROFILE
SEE PROFILE

All content following this page was uploaded by Subodh Kumar Datta Datta on 20 July 2021.

The user has requested enhancement of the downloaded file.

LS - An International Journal of Life Sciences
DOI: 10.5958/2319-1198.2020.00017.2 IndianJournals.com
A product of Diva Enterprises Pvt. Ltd.

Genetic Diversity and Improvement of Bougainvillea

S.K. Datta
Retired Scientist, CSIR-National Botanical Research Institute, Lucknow; Ex CSIR Emeritus scientist, Bose Institute,
Kolkata; Ex DBT-NER Visiting Research Professor, Assam University, India
Email id: [email protected]

Received: 10-03-2021; Accepted: 30-05-2021

ABSTRACT
Bougainvillea is a very important floriculture crop for multipurpose use. A good number of varieties
have been developed in different parts of world/India through detection and isolation of bud sports,
hybridization and induced mutation. All basic scientific information are available on different aspects
of Bougainvillea. Considering its importance more work should be initiated for development of
improved varieties through careful selective breeding. Conservation of genetic diversity of
Bougainvillea is an important aspect. Breeding work may be initiated to develop more and more
strains tolerant to climate change.

Keywords: Bougainvillea, Elemental species, Bud sport, Genetic diversity

INTRODUCTION importance with very showy and colourful bracts

(Heimerl, 1900). All the present day Bougainvillea
Bougainvillea is one of the most important and popular
cultivars being grown in India are either exotics introduced
perennial ornamental crop in floriculture for multipurpose
from abroad (Africa, USA, U.K., South Africa, Australia,
use and a very suitable plant for multidisciplinary research
Philippines, Sri Lanka, Jamaica, Canary Island, Singapore,
work due to its numerous bract colours, leaf characters
Java, West Indies etc.) or developed by concentrated
and plant stature. Fascinating variegation in leaves is added
beauty for its attraction. It is a versatile plant which grows efforts of nurserymen, individuals or scientific institutions.
very well under the tropical and subtropical conditions. Around the 1930s, when the three species were grown
The plant grows well in various climatological zones and together, many hybrid crosses were created almost
also in almost all types of soil. Considering its present spontaneously in East Africa, India, the Canary Islands,
status, it is necessary to review the genetic diversity and Australia, North America, and the Philippines.
its improvement for future research on Bougainvillea. Literature on description of elemental species and mode
Bougainvillea belongs to the family Nyctaginaceae and it of origin, sports (cultivated species), hybrids and mutants
is the only genus in the monotypic subtribe are insufficient and all the literature are not available at
Bougainvilleineae (Gills, 1976) and has originated in one place. The classification and nomenclature of the
tropical and sub-tropical South America. It was found species, varieties or cultivars of Bougainvillea are not very
growing wild besides being grown as a cultivated garden much well understood and still there are many
plant in Brazil, Colombia, Ecuador and Peru. Out of 10 discrepancies in the views of various workers. In India,
species, three (B. spectabilis Wild., B. glabra Choisy and collection of Bougainvillea and development of new
B. peruviana Humb. and Bonp.) are of horticultural varieties was started by amateurs, commercial or non-

Volume 10, Number 2, May-August, 2021, pp. 61-79 61

 S.K. Datta

commercial professionals, researchers, nurserymen etc. make the environment colourful and magnificent. Division
Appreciable knowledge has been built-up on bougainvillea of Floriculture and Landscaping, Indian Agricultural
breeding and commercial features have been identified in research Institute, New Delhi was recognized as the
different elemental species and cultivars. Breeders and International Registration Authority in 1966 for
amateur growers reported noticeable concepts on Bougainvillea cultivars. Since then the Division is
breeding. The review article will focus on important compiling, maintaining, publishing lists of names of
aspects of bougainvillea breeding and related background cultivars and registering new varieties. Voluminous
knowledge generated by breeders. Bougainvillea now compilation work resulted publication of first “The
stands as a very important floriculture crop for its International Bougainvillea Check List” in 1981 and
multipurpose use. Attempts have been made to gather subsequently “The New International Bougainvillea Check
scattered knowledge and breeding technology and to utilize List” in 1999 (Singh et al., 1999). These checklists are
such wealth of information for future breeding of immense importance to Bougainvillea lovers,
programmes to develop varieties as per need base and researchers, floriculturists, landscape architects, town
societal benefit. planners, gardeners of the world. Most of the present
day varieties have arisen through bud sports, chance
The article covers genetic diversity and their mode of
crossing and induced mutation. India is one of the major
origin and improvement through breeding. Range of repository of wide range of genotypes of Bougainvillea
diversity in different characters, source of diversification and is perhaps the foremost among nations which have
and different approaches for further improvement have done appreciable good work on Bougainvillea. Voluminous
been highlighted step by step. Attempt has been made to work has been done in India on different aspects like
put together all information to develop a complete enrichment of germplasm, agro-technology,
documentation of the results of the research conducted characterization and improvement at Indian Agricultural
by different organizations. The article gives a systematic research institute (IARI), New Delhi; National Botanical
scenario on bougainvillea research in India in a nutshell. Research Institute (NBRI), Lucknow; Indian Institute of
The report has been prepared only from published data/ Horticultural Research (IIHR), Bangalore; Lalbaugh
knowledge as a review document. The early history of Gardens, Bangalore; Bhaba Atomic Research Centre
bougainvillea regarding its origin, nomenclature, (BARC), Trombay; Agri-Horticultural Society, Calcutta.
distribution etc. have been recently compiled (Datta et Different Agriculture Universities, amateur growers and
al., 2017). nurserymen are also maintaining good amount of cultivars
due to its increasing demand and popularity. Bougainvillea
Introduction in India profusely flowers almost all the year round except in rainy
Bougainvilleas are widely grown in Indian gardens. The season and very severe winter.
Bougainvillea spectabilis was the first to be introduced
in India from Europe in 1860 followed by B. glabra cv. Varieties Reported from India
‘Splendens’ (1969), B. x buttiana cvs. ‘Scarlet Queen’ It is little difficult to mention the exact number of total
(1920) and ‘Mrs. Butt’ (1923) and B. peruviana cv. varieties available in India. Different workers have
‘Princes Margaret Rose’ in 1935. Most of these reported the names of varieties available with them. There
Bougainvilleas were introduced in Calcutta by the then are duplications in name of varieties. There are more than
Royal Agri-Horticultural Society of India. Later the 400 varieties growing in the Indian gardens and following
introductions were made by the Agri-Horticultural Society, are the names which call attention to popular varieties -
Madras and the Lalbaugh Botanic Gardens, Bangalore. ‘Abraham Kavoor’, ‘Abhimanyu’, ‘Aida’, ‘Aida Variegata’,
Since then it is very much popular in nearly every city, ‘Alba’, ‘Alick Lancaster’, ‘Alizon Davy’, ‘Amaranth’,
town or rural hamlet across the country. Bugainvillea is a ‘Amarnath’ ‘Amarault’, ‘Anindita’, ‘Arjuna’, ‘Annabella’,
very versatile plant posses various colour and bloom to ‘Archana’, ‘Aruna’, ‘Arunima’, ‘Asia’, ‘Autumn’, ‘Baby

62 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

Margaret rose’, ‘Bangalore Variegata’, ‘Barbara Karst’, ‘Magenta Queen’, ‘Mahara’, ‘Mahara Purple’, ‘Mahara
‘Beauty’, ‘B. A. Rama Rao’, ‘B.G.K. Hamilton’, ‘Begum Variegata’, ‘Mahara Variegata Abnormal Leaves’, ‘Mahara
Sikander’, ‘Baron Hado’, ‘Begum Ali Yawar Jung’, ‘Dr. White’, ‘Maharaja of Mysore’, ‘Mahatma Gandhi’,
Bhabha’, ‘Bonfire’, ‘Brenda Ohare’, ‘Brilliant’, ‘Blondie’, ‘Mahatma Gandhi Variegated’, ‘Maharaja of Mysore’,
‘Bois de Rose’, ‘Brilliance’, ‘Brasiliensis’, ‘Buddhadas’, ‘Manohar Chandra Variegated’, ‘Marigowda’, ‘Margery
‘Camarillo Fiesta’, ‘Carlton Corea’, ‘Cascade’, ‘Cherry Lloyd’, ‘Manohar Chandra’, ‘Mary Palmer Special’,
Blossom’, ‘Cherry Ripe’, ‘Chandrabieri’, ‘Charles ‘Mataji Agnihotri’, ‘Mauve Queen’, ‘Meera’, ‘Meera
Williams’, ‘Cleopatra’, ‘Chinese Cracker’, ‘Chitra’, Sport’, ‘Midget’, ‘Minyatab’, ‘Meriol Fitzpatrick’, ‘Mrs.
‘Chitravati’, ‘Coral Dawn’, ‘Common Rose’, ‘Conquest’, Bakery’, ‘Mary Palmer’, ‘Miss Manila Red’, ‘Maud
‘Crimson King’, ‘Crimson Lake’, ‘Crispa’, ‘Cypheri’, Chattleburgh’, ‘Mrs. Butt Magenta’, ‘Mrs. Butt Scarlet’,
‘Daya’, ‘Deep Cherry’, ‘Dogstar ’, ‘Dorothy ‘Mrs. Chico’, ‘Mrs. Eva’, ‘Mrs. Eva Variegata’, ‘Meriol
Jivarajadasa’, ‘Double Delight’, ‘Dr. Hado’, ‘Dream’, Fitzpatrick’, ‘Mrs. Delux Perry’, ‘Mrs. Fraser’, ‘Mrs.
‘Durga’s Delight’, ‘Dwarf Gem’, ‘Dr. H.B. Singh’, ‘Dr. G.S. Randhwa’, ‘Mrs. Kay Malvenan’, ‘Mrs. Lancaster’,
B.P. Pal’, ‘Dr. P.V. Sane’, ‘Easter Parade’, ‘Eclipse’, ‘Mrs. Marie Buck’, ‘Mrs. Butt’, ‘Mrs. H. C. Buck’, ‘Mrs.
‘Ecstasy’, ‘Eisha’, ‘Elizabeth’, ‘Elizabeth Angus’, ‘Ena R. B. Carrick’, ‘Mrs. McClean’, ‘Mrs. McClean Nirmal’,
Heynekar’, ‘Dr. R.R. Pal’, ‘Enid Lancaster’, ‘Enid ‘Mundanna’, ‘Natalii’, ‘Mrs. Oliver Perry’, ‘Mrs. R.B.
Walker’, ‘Fantasy’, ‘Formosa’, ‘Easter Parade’, ‘Ethiraj’, Carrick’, ‘Mudanna’, ‘Nawab Ali Yawar Jung’, ‘New
‘Eusga’, ‘Exquisite’, ‘Feathery Fantasy’, ‘Feathery Red’, ‘Nirmal’, ‘Nirmal Chandra’, ‘New Red’, ‘Nigrette’,
Fantasy Bi Coloured’, ‘Filoman’, ‘Flame’, ‘Floribunda’, ‘Orange Glory’, ‘Orange Sceptre’, ‘Odisee’, ‘Orange
‘Frazer ’ ‘Fuschea’, ‘Gagarin’, ‘Gangamma’, King’, ‘Palekar’, ‘Padma’, ‘Padmi’, ‘Padmini’, ‘Pallavi’,
‘Gangaswamy’, ‘Garnet Glory’, ‘Gem’, ‘Geoffrey ‘Pearl’, ‘Penang’, ‘Partha’, ‘Parthasarathy’, ‘Pink Beauty’,
Nagpal’, ‘Gillian Greensmith’, ‘Glabra Magnifica’, ‘Glabra ‘Pisil’, ‘Perfection’, ‘Philips’, ‘Philoman’, ‘Poultoni
Magnifica Trailii’, ‘Glabra’, ‘Glady’s Hepbum’, ‘Glabar Special’, ‘Pixie’, ‘Pixie Variegata’, ‘Poultoni’, ‘Poultoni
var Cypheri’, ‘Gloriosus’, ‘Glabra Sanderiana’, ‘Glabra Variegata’, ‘President Roosevelt’, ‘Pradhan’s Pink’,
Variegata’, ‘Golden Glow’, ‘Golden Giant’, ‘Golden ‘Pradhan’s Profusion’, ‘Preeti’, ‘President’, ‘Prolific’,
Queen’, ‘Gopal Swamy’, ‘Gopal’, ‘Gokul’, ‘Gulaby’, ‘Princess Margaret Rose’, ‘Profusion’, ‘Purple King’,
‘H.C. Buck’, ‘Happiness’, ‘Happikiness’, ‘Hawaiian ‘Purple Mrs. Butt’, ‘Purple Prince’, ‘Purple Star’, ‘Purple
White’, ‘Hawaiin Beauty’, ‘Himani’, ‘Hiawatha’, Gem’, ‘Purple Queen’, ‘Purple Rose’, ‘Purple Wonder’,
‘Intermedia’, ‘Issabel Green Smith’, ‘Isobel’, ‘Jane ‘Queen Elizabeth’, ‘R.S. Bhutt’, ‘Rani’, ‘Raman’, ‘Ratna’,
Stanfeld’, ‘Jasper Rose’, ‘Jawaharlal Nehru’, ‘Jaya’, ‘Red Glory’, ‘Radha’, ‘Rambha’, ‘Ranee’, ‘Rhodamine’,
‘Jayalakshmi’, ‘Jayalakshmi Variegata’, ‘Jennifer’, ‘Red Glory Improved’, ‘Red September’, ‘Red Triangle’,
‘Jennifer Fernie’, ‘Jennifer Nagpal’, ‘Joe’, ‘Joe de Livera’, ‘Refulgens’, ‘Rosea’, ‘Rosa Multiflora’, ‘Rose Queen’,
‘John Latin’, ‘Jubilee’, ‘Kayata’, ‘Kalyani’, ‘Killie ‘Ruarka’, ‘Rosea Fuchsea’, ‘Roseville’s Delight’, ‘Scarlet
Campbell’, ‘Krumbeigel’, ‘Kuvempu’, ‘L.N. Birla’, ‘Lady O’Hara’, ‘Sachidananda’, ‘Scarlet Glory’, ‘Sensation’,
Elizabeth’, ‘Lady Hope’, ‘Lady Hudson of Ceylon’, ‘Lady ‘Scarlet Queen’, ‘Scarlet Queen Variegated’, ‘Sharma’,
Mary Baring’, ‘Lady Mountbatten’, ‘Lady Richard’, ‘Sholay’, ‘Simon Anselm’, ‘Shubhra’, ‘Shweta’, ‘Silver
‘Lalbagh’, ‘Lal Baugh Louis Wathen’ ‘Lateritia’, ‘Lavender Top’, ‘Snow Queen’, ‘Soundarya’, ‘Sova’, ‘Spring
Glory’, ‘Lavender Queen’, ‘Lakshmi’, ‘Laxminarayana’, Festival’, ‘Singapore Red’, ‘Singapore Dark Red’,
‘Lazat of Mysore’, ‘Leah Nagpal’, ‘Leopaldi’, ‘Lilac ‘Sonnet’, ‘Srinivasa’, ‘Speciosa’, ‘Spectabilis’,
Beauty’, ‘Lilac Puff’, ‘Lilac Queen’, ‘Liliac Perfection’, ‘Spectabilis Variegated’, ‘Specaible’, ‘Spitfire’,
‘Lilacina’, ‘Limousine’, ‘Lord Willingdon’, ‘Los Banos ‘Splendens’, ‘Sumatra’, ‘Sri Durga’, ‘Srimati Biezli’,
Beauty’, ‘Los Banos Variegata’, ‘Lucifer Red’, ‘Louise ‘Stanza’, ‘Star’, ‘Star Mauve’, ‘Superb’, ‘Summer Time’,
Wathen’, ‘Louise Wathen Intermediate’, ‘Louise Wathen ‘Sundari’, ‘Sweet Heart’, ‘Surekha’, ‘Suverna’, ‘Tetra
Variegated’, ‘Louise Wathen Mediopicta’, ‘Machakos’, Mrs. McClean’, ‘Sydney’, ‘Temple Fire’, ‘Tetra

Volume 10, Number 2, May-August, 2021 63

 S.K. Datta

McClean’, ‘Thimma’, ‘Texas Dawn’, ‘Theresa Jacobs’, Baring’, ‘Louise Wathen Medipicta’ (sy. ‘Louise Wathen
‘Trailli’, ‘Triangle’, ‘Torch Glow’, ‘Trinidad’, ‘Tomato variegata’), ‘Louise Wathen’ (syn. Orange Glory),
Red’, ‘Tyrian Rose’, ‘Udai Chandra’, ‘Usha’, ‘Vellayani’, ‘Magenta Queen’ (syn. ‘Mrs Butt Magenta’), ‘Mary
‘Turley’s Special’, ‘Versicolour’, ‘Vijaylakshmi’, ‘Verna Baring’, ‘Purple King’, ‘Purple Queen’, ‘Mrs Butt
Nagpal’, ‘Vijaya’, ‘Vithal Variegated’, ‘Vishakha’, Variegata’, ‘Mrs McClean’ (syn ‘Orange King’ Practorius),
‘Vesuvius’, ‘Vithal’, ‘Wajid Ali Shah’, ‘Walker’, ‘Win ‘Purple Prince’, ‘Roseville’s Delight’ (syn ‘Dona Rosita
Some’, ‘Yellow Queen’, ‘Zakir Hussain’, ‘Zakiriana’, Delight’, ‘Doubloom’), ‘Rao’, ‘Scarlet Queen’, ‘Surekha’,
‘Zakiriana Variegate’, ‘Zulu Queen’, ‘Zinna Barat’. ‘Scarlet Queen Variegata’, ‘Versicolour’, ‘Vellayani’
‘Yellow Queen’, etc. Description of elemental species and
Diversity important role played by Bougainvillea cv. ‘Mrs. Butt’ for
the evolution of many important cultivar of modern
Natural hybridization and sports were common
bougainvillea have been reported in some early publications
occurrences all over the world to create diversification.
(Holttum, 1955a,b, 1957; Pancho and Bardenas, 1959;
Bougainvillea is most important and widely accepted in
Marigowada, 1961; Sharma, 1962 and Zadoo et al.,
floriculture due to wide range of variability. Diversification
1976).
resulted due to geographical movement of elemental
species, adaptation, selection pressure, spontaneous
Diversity in Use
mutation, natural and intentional breeding, induced
mutation etc. Spontaneous hybridization and/or bud sports Bougainvilleas is a multipurpose plant due to the variation
specially created wide range of variations in bract shape, in habit, growth and colour pattern of different cultivars.
size and colour during cultivation. Diversity was created due to selection pressure and now
Bougainvilleas are grown as bush, climbers, specimen
Diversity in morphological and horticultural aspects of
plant, hedges, topiaries, ground covers, standards, on
three species and 40 cultivars were analysed and suggested
pergolas and trees, arches, pot culture, cut flowers, slopes
that the diversity may be primarily due to variability within
and mound, bonsai, hanging basket, cascade etc. It is
species B. spectabilis and B. glabra through indiscriminate
also a very important plant for landscaping. It is drought,
hybridization and somatic mutations among themselves
saline and pollution tolerant and can be planted in the
and with B. peruviana (Zadoo et al., 1975a). The salient
industrial regions and in the areas where other ornamental
morphological feature of four important species
flowering plants do not easily thrive well. Genotypes
(Bougainvillea spectabilis, Bougainvillea peruviana,
suitable for each category have been selected (Jana and
Bougainvillea glabra and ‘Mrs Butt’) have been studied
Maithi, 1986; Singh, 1996; Datta et al., 2017; Jayanthi,
(Kobayashi et al., 2007; Pancho and Bardenas, 1959).
2000; Kumar et al., 2007; Kumar et al., 2015; Sindhu,
‘Mrs. Butt’ is a natural hybrid between B. peruviana and
2001).
B. glabra. It was first recorded by (Mrs. R.V. Butt
Holttum, 1955a, b). Mrs Butt came to India in 1923 from
Diversity in Morphological Characters
Kew to Bangalore. ‘Mrs Butt’ is having the characters of
both parents i.e. leaves like B. peruviana and floral bracts Bougainvillea bracts are available in almost all colour shades
like B. glabra (Holttum, 1955b). Spontaneous mutations like red, magenta, mauve, purple, orange, yellow, white
have played very important role in developing a number etc. Bracts are also available in bicolours and multicolor
of new bract colours like dianthus purple, phlox purple, form. On the basis of bract nature, Bougainvillea is divided
burnt orange, spanish orange, majolica yellow etc. from into (1) Single bracted and (2) Double or Multi-bracted
‘Mrs Butt’. Some important varieties which have either Bougainvillea. Variegated foliage bougainvilleas are
direct or indirect link with Mrs Butt are ‘Alick Lancaster’, sometimes recommended as third group. But in reality,
‘Archana’, ‘Bhabha’, ‘Enid Lancaster’, ‘Gangaswamy’, all variegated varieties are developed either through
‘Gangamma’, ‘Golden Glow’, ‘Kuvempu’, ‘Lady Mary spontaneous or induced mutations from existing single

64 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

or double bracted varieties. Majority of Bougainvillea chemical and biochemical, DNA markers have been taken
belong to single bracted group with wide range of bract into consideration. Zhou et al. (2011) characterized 65
colour spectrum. The wide range of adaptability along introduced varieties of bougainvillea to avoid confused
with diverging habitate and broad spectrum of bract colour problems in synonyms and homonym of bougainvillea in
and variegated foliage place this plant on a very important China. They established the classification system of these
position among ornamentals. Duble bracted group of varieties based on the “International Code of Nomenclature
Bougainvillea consists of only four normal varieties for Cultivated Plant”. Extensive work has been carried
(‘Cherry Blossom’, ‘Los Banos Beauty’, ‘Mahara’ and out at CSIR-NBRI, Lucknow for morphological, horto-
‘Roseville’s Delight’). taxonomical, bio-chemical and cytogenetical
characterization of different varieties to estimate the
Characterization : Horto-Taxonomy diversity in different characters. Under characterization
programme different characters like habit; growth; bract
Early reports are always pathfinders for future research.
colour and size; flowering behavior; flower size; leaf colour
Earlier studies at CSIR-NBRI on characterization of
and size; branch and leaf number; leaf shape; spine number
bougainvillea is so important that it should be highlighted
and size; stem colour; stomatal index; moisture content
for researchers. Horto-taxonomic description of the
of bract; pollen grain sterility and ornamentation;
leading Bougainvillea species/cultivars have been prepared
chlorophyll content of leaves; bract pigment and phenolic
according to their affinities to the species and hybrid
compounds in leaves and bracts have been studied in detail
seedlings (Sharma, 1986; 1996; Khoshoo, 1971a, b; Zadoo
in different varieties. Few reports are also available from
et al., 1975a; Jayanthi, 1999). Many workers have
other institutions/universities on characterization to
characterized bougainvillea varieties from time to time to
estimate diversity (Jayanthi, 1999; Sharma, 1996; Chandel
understand the diversity. Many cultivars have developed
et al., 2015; Kumar, 2001; Kumar et al., 2007; López and
as a result of interbreeding among different species and
Galetto, 2002; Jayanthi, 2000; Kumar, 2001; Singh, 1996;
cultivars. Induced and spontaneous mutations and
Sindhu, 2001; Zadoo et al., 1975a; Zhou et al., 2011)
chromosomal manipulations also played an important role
in developing varieties. Some of these varieties are
Bract Colour
intermediate types that are more like one parent and some
are like other. Many hybridizers in different countries Bract colour plays a significant role in identifying/
introduce and name seedlings without evaluation or beautifying different cultivars and makes Bougainvillea a
registration. Lack of proper pedigree records and other noticeable/wonderful colourful garden plant. On the basis
relevant data before the cultivar is named or included in of Royal Horticultural Society of London all single bracted
further breeding programme, not only creates confusion bougainvillea have been classified into eight colour groups
but gives rise to a large number of cultivars of uncertain on the basis of colour variance viz. Red-Purple Group,
identity which would often not merit separate taxonomic Purple-Violet Group, Purple Group, Red Group, Violet
status and entity. Many times the same cultivar has been Group, Orange and Orange-Red Group, White Group,
differently named at different places and in some cases Bi-coloured Group. In some varieties there was change
even the same name has been given to the cultivars of in colour shade of bract with age. In some cases the
different origin. Characterization of germplasm is very colour was mixed. Out of eight multi-bracted group,
important for estimating/understanding genetic diversity, colour of matured bracts of ‘Cherry Blossom’, ‘Mahara’
for proper identification, detection of phylogenetic and ‘Mahara Variegata’ were Red-Purple Group, ‘Los
relationship, taxonomic status, preparation of catalogue, Banos Beauty’ was Purple Group and ‘Los Banos
identification of desirable/novel genes, hybridization, Variegata’ was Purple-Violet Group, ‘Archana’, ‘Pallavi’
registration, plant variety protection, farmer’s right etc. and ‘Roseville’s Delight’ were bi-coloured. Some
Different parameters of cytology, morphology, physiology, bougainvillea varieties changes their bract colour with age.

Volume 10, Number 2, May-August, 2021 65

 S.K. Datta

Bract Size : Size of bract varied from variety to variety Shape of the Leaf Apex
and these variations were observed in each colour group.
Leaf apex of all the varieties were categorized under five
Single bracted – bract length 1.75 -5.24 cm and width
groups i.e., acute, acuminate, acute to mucronate, acute
1.07-3.92 cm; Multibracted – bract length 0.49-2.82 cm
to acuminate and mucronate. Shape of leaf tip varied
and width 0.28-1.87cm.
among the varieties in each colour group.
Flower Size i.e. the length of flower tube in single bracted
varieties are 0.57-2.40 cm and diameter of the star are Number of Branch
0.43-1.00cm. In double bracted varieties there is no
flower. Number of branches and leaves per 30 cm mature shoots
varied in different cultivars. In single bracted cultivars
Inflorescence cyme either born on main shoot (B. shoot number varied from 3.80-20.20 and leaf number
peruviana and B. x buttiana group) or on lateral branches 24.20-198.40 whereas branches were 5.00-10.00 and
(B. glabra, B. spectabilis, specto-peruviana and spectro- leaves were 43.00-284.00 in multibracted cultivars.
glabra group). The cyme may divide once or twice in B.
spectabilis and B. glabra, while in B. peruviana, which Number of Thorn
has unbranched shoot it divides more than twice. Flower
tubes varies in shape and colour in basal species. Different Number per 30 cm length of mature stem were 6.00-
cultivars show various types of tube shape and grade of 28.60 in single bracted and 6.20-14.20 in multibracted
pubescence which in some cultivars of B. x buttiana is cultivars.
imperfectly developed. Tube lacks the conspicuous star,
filament length is reduced while style remains unchanged. Thorn size (length)
With the results anthers are arranged in a bunch at the
In single bracted varieties it varied from 0.34-2.48 and
base of protruding style.
0.30-1.72 in multibracted cultivars. Different cultivars
have been categorized on the basis of thorn character
Leaf Colour
(Sindhu and Swaroop, 2015) - Thorn less cultivars,
Colour of leaves of all the varieties were noted at three Cultivars with few thorn and Cultivars with vigorous
stages (new leaf initiation stage, young and matured). thorns.
Leaves of all the varieties were categorized into different
groups viz., Yellow-Green Group, Green Group, Brown Stem Colour
Group, Greyed-Orange Group, Variegated, Greyed-Purple,
Coppery-Red, Red-Purple, Greenish-Brown and Greyed- The colour of mature stem of all the varieties were found
Brown on the basis of their colour similarity to colour to be either Brown Group, Grey group, Greyed-Green or
chart. In most of the group the colour of leaves changed Greyed-Brown Group. Shades of each group varied from
from new leaf to matured leaf. variety to variety.

Leaf Size Stomatal Index

A wide range of variability in leaf length, leaf width and Variation in number per unit area and size of stomata and
petiole length was observed among the varieties of each number of chloroplasts in guard cells were recorded both
group. The varieties were categorized into different among the varieties of each group and also between the
classes on the basis of leaf size. Size of single bracted varieties of different groups. In single bracted cultivars
leaf- length 3.06-8.60 cm, width 2.01-6.34 cm and length stomata size were 32.22-40.00 µm (length) and 23.06-
of petiole 0.08-3.57 cm. Multibracted leaf length 4.31- 30.56 µm (width). In multibracted length and width were
7.73 cm, width 2.81-6.22 cm and length of petiole 0.87- 38.61-40.00 µm and 25.75-29.44 µm respectively.
1.93 cm. Number of stomata were 17.25-51.40 and chloroplast

66 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

number per guard cell were 13.50-26.20 in single bracted categorized into 12 major clusters and characters may be
cultivars. Similarly the same in multibracted were 16.25- used for identification of cultivars.
19.87 and 15.50-25.20, respectively.
DIVERSITY IN CYTOGENETICS
Pollen Grain Size and sterility
Bougainvillea is a very interesting material for cytogenetical
Pollen grain size (polar diameter) of all the varieties were studies to trace out their relationship with wild ancestors
uniform i.e., ‘Normal’ (almost same size). ‘Small’ and like other ornamentals. Bougainvillea shows a great deal
‘Big’ sized pollen grains in addition to ‘Normal’ pollen of cytological polymorphism. CSIR-NBRI, Lucknow
grains were recorded in some of the varieties. The smallest made concentrated efforts to understand the genetic and
pollen grain of the Big category was larger than the largest cytogenetic basis for the diversity in bougainvillea. This
pollen grain of the normal category and the largest grain basic studies helped to prepare a conscious breeding
of the small category was smaller than the smallest grain methodology for further genetic improvement. All the
of the normal category. Pollen grain size varied from 24.46- present day garden forms of bougainvillea are of hybrid
48.21 µm and sterility 13.55-99.04 per cent. Pollen grain origin raised over a number of years and named differently
ornamentation characters are genetically controlled. In by different workers and nurserymen. Cytogenetic studies
course of time a wide range of variability have been were concentrated precisely on basic aspects like
developed in ornamentation pattern. Pollen grains of determination of chromosome number, mitotic and meiotic
Bougainvillea are 3(-4) colpate, shape is prolate spheroidal systems, analysis of karyotype, DNA content etc. Studies
to oblate spheroidal and exine is prominently reticulate. covered three basal Bougainvillea species, hybrid groups,
Two sets of bacula are present in most cultivars. The triploid hybrid cultivars and induced tetraploids. Efforts
outer bigger set forms the muri and the inner set supports were also made to understand the pollination mechanism,
the lumina. The shape and size of the lumina are variable breeding system and cross compatibility relationship. Such
among the cultivars. Pollen morphology of three studies opened a new way to manipulate chromosomes
elementary sps. B. glabra, B. spectabilis and B. peruviana for synthesizing new variety. Cytological studies at CSIR-
and cultivars derived from them have been studied. NBRI played very significant role for improvement of
Variations in pollen morphology may be very helpful in bougainvillea (Datta and Banerji, 1995; Khoshoo, 1968,
understanding the interrelationship among the varieties. 1986, 1990; Ohri, 1995; Zadoo et al., 1975a, b, c, d, e, f;
Zadoo and Khoshoo, 1975; Ohri and Zadoo, 1975, 1979,
Varietal differences in relation to exine ornamentation,
1986; Zadoo et al., 1976; Xu et al., 2009; Ninan et al.,
coupled with shape of the grain is reported where one
1959).
type contained smaller brochi and ± spheroidal shape and
the other type having larger brochi and prolate spheroidal Chromosome number was determined by different
to subprolate in shape (Erdtman, 1952; Nair, 1961). Pollen workers in different varieties like 2n=20 in B. glabra,
grains of different species were dissimilar in the shape of 2n=34 in different species and hybrid (Wilson, 1947),
apex of brochi besides differences in pollen size and 2n=34, 2n=20 in different cultivars of B. glabra and B.
number and size of brochi and sterility (Swarup and Singh, spectabilis (Ninan et al., 1959), 2n=32 in two varieties
1964; Ninan et al., 1959; Pramanick et al., 2015). Pollen resembled to B. spectabilis (Sen and Sen, 1954), 2n=34
ornamentation patterns provide useful characters for in 37 varieties and 2n=51 in three varieties (Sharma and
identification of the species, B. spectabilis (Lopez and Bhattacharya, 1960). Banerji and Banda (1967) reported
Galetto, 2002; Takashashi and Skvaria, 1991; Pramanick natural triploidy and Begum and Datta (1971) reported
et al., 2015). Kumar et al. (2015) selected 38 new ‘n’ number (n=8). Different types of meiotic
morphological descriptors as per UPOV guidelines and abnormalities viz. lagging, early and late separation,
analysed hundred cultivars and detected a significant unequal separation, inversion heterozygotes, haploidy etc.
amount of diversity. Bougainvillea cultivars have been have been reported in some cultivars ‘Gangamma,

Volume 10, Number 2, May-August, 2021 67

 S.K. Datta

‘Gangaswamy, ‘Thimma’ and ‘Velliyani’ (Sen and Sen, number of varieties belonging to B. glabra, B. buttiana
1954; Sharma and Bhattacharya, 1960; Pancho and and B. spectabillis were studied and 2n=34 chromosomes
Capinpin, 1961). were recorded. However, in some buds from red sectorial
part of ‘Marry Palmer’ 2n=20 chromosomes were
Studies on pollination mechanism, breeding system and
detected (Ninan et al., 1959).
cross compatibility in different elemental species and
‘Mrs. H.C. Buck’ and its family showed that 80 percent Cytogenetical, pollination mechanism and cross
of garden Bougainvillea are male and female sterile. It compatibility studies on Bougainvillea at CSIR-NBRI
was interesting that fertile forms do not set seed on selfing clearly indicated that choice of the female parents is limited
but crossing readily set seed. Hybrid development was in any breeding programme. This really encroaches upon
identical to ‘Mrs. H.C. Buck’ as tested by morphological noticeable restriction in the choice of parents and excludes
and biochemical characters. All the cultivars of three cultivars that may be sterile but otherwise progress is
elemental species of Bougainvillea are diploid (2n=34) and possible unless methods are evolved for restoring pollen
while B. glabra and B. spectabilis show normal 17 and/or seed fertility in order to have a broad based
bivalents, B. peruviana is an interchange heterozygote germplasm. NBRI made a very interesting study and
with regular 15 bivalents and a ring of four (Zadoo et al., opened a new dimension for Bougainvillea breeding
1975b). Zadoo et al. (1975c, f) studied 70 cultivars through induction of tetraploidy and restoration of fertility
belonging to three species and hybrid groups and reported in sterile cultivars. Cultivars belonging to the three hybrid
2n=34 (diploid) chromosomes and 80 to 90 per cent pollen groups are sterile but they possess a number of desirable
fertility in all the taxa except ‘Perfection’ and ‘Poultonii characters for breeding. 80% of the cultivars are pollen
Special’ which were triploid (2n=51). Induced tetraploidy and seed sterile while only those belonging to the elemental
restored fertility in sterile cultivars. Induced tetraploids species and triploids are fertile. The major biggest
of the fertile diploid cultivars are self-incompatible, but constraint on further breeding and improvement of
set seeds readily on crossing. Irregular meiosis was bougainvillea is the narrow range of germplasm availability.
observed in diploid progenitors and tetraploid counterparts ‘Tetra Mrs. McClean” (1969) is such a synthesized
showed predominant bivalent pairing. These studies helped tetraploid variety. Tetraploidy (2n=68) has been induced
to introduce a new perspective in bougainvillea breeding. in a number of cultivars i.e. ‘Shubhra’, ‘Mrs. McCleans’,
Analysis of 60 cultivars showed 2n=34 (diploid) ‘President Roosdevelt’, ‘Lady Mary Baring’, Zakiriana’,
chromosomes in 58 and two were triploid (2n=51) and ‘Thimma’, etc. A breeder can therefore broaden the
all were self sterile. Ten cultivars have pollen stainability germplasm base by inducing tetraploidy in desirable, highly
ranging from 50 to 90 percent, out of which seven have floriferous and colourful cultivars. These can be further
50 to 80 percent ovule fertility. DNA content of B. utilized in planned hybridization programmes to produce
peruviana was found to be significantly different from polyploidy and aneuploid hybrids with entirely novel
B. spectabilis and B. glabra. DNA content of three hybrid horticultural attributes (Zadoo et al., 1975e; Khoshoo and
group i.e. B. specto-glabra, B. specto peruviana, B. Zadoo, 1969; Zadoo et al., 1975c).
glabra peruviana and synthetic hybrid were intermediate
between parents and they found direct correlation of DNA DIVERSITY IN CHEMICAL CHARACTERS
content and ploidy level (Khoshoo and Zadoo, 1969; Ohri
Phenolic Compounds in Leaves
and Khoshoo, 1982; Zadoo et al., 1975c). Cytology,
cytokinesis and microspore formation were studied on Phenolic compounds (flavonol, flavones, anthocyanins,
13 variegated cultivars and detected gametic chromosome coumarins and others) have been studied in leaves and
number n=17. All variegated cultivars were grouped into bracts by TLC methods and used as a tool for
three categories on the basis of degree of meiotic chemotaxonomical characterization. Distribution of some
abnormality (Srinivas and Bhat, 1982). Natural triploidy of these compounds varies not only in different species
has been reported by Banerji and Banda (1967). A large but also in cultivars of the same species. Inter and Intra

68 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

varietal similarities were observed on the basis of 6-O--glucoside and various derivatives of bougainvillein-
distribution of phenolic compounds in leaves (Nageshwar v (betanidin 6-O--sophoroside), i.e. mono- and
and Radhakrishnaiah, 1994; Jayanthi, 1999). diglucosylsophorosides which are acylated with 4-
coumaric and caffeic acid (mono- and diesters). Besides
Phenolic Compounds in Bracts the betacyanins, B. glabra bracts accumulated large
amounts of flavonols (kaempferol and quercetin
Phenolic compounds in bracts of different varieties were
conjugates), reaching ratios of flavonol to betacyanin of
determined by TLC methods the same thin layer
1:1. ‘Marry Palmer’ with white shoot sported from ‘Mrs.
chromatographic methods as done in case of leaves. A
H.C. Buck’ with magenta bract. The magenta betacyanins
total of 17 spots were detected among all the varieties
are in some way suppressed and it leads to white bracts
tested here. Inter-and intra-varietal similarities were
being flushed with a mild pink tint (c.f. Crane and
observed on the basis of distribution of phenolic
Lawrence, 1952). In ‘Marry Palmer’ bracts in a single
compounds in bracts (Jayanthi, 1999).
plant are either magenta or parchment white, or even half
magenta and half white creating an outstanding novelty.
Bract Pigment
Such blotched bracts are seen in several hybrids involving
Inter varietal differences were observed and confirmed ‘Shubhra’. Variations in bract colour with variation of
by number of peaks and their O.D. values (Jayanthi, 1999). environment (cool and warm) have been recorded in
‘Marry Palmer’ and hybrid between ‘Shubhra’ and
DIVERSITY IN BRACT PIGMENT ‘Jennifer’. No conclusive reports are available about the
basis of underlying colour change in sport, hybrid and
Although there are a wide range of variations in bract mutant cultivars.
colours in Bougainvillea, no systematic work has been
done on genetic and/or chemical basis of bract pigment
Diversity in Extract use
composition. Diversity in pigments have been reported
from morphological variations among different cultivars, Bract, leaf, root and plant extracts can be used - as - acid
hybrids, sports, mutants and few experimental analysis. base indicator in titration, anthelmintic activity, anti-fungal,
The red and other colours in Bougainvillea are due to anti ulcer, antimicrobial activities, anti-diabetic, anti-
betacyanin which are not related to anthocyanins, but are diarrhoeal, antifertility potential, anti-inflammatory activity,
similar to auxins. Yellow pigments are known as insecticidal, in vitro anti-bacterial, cardiotonic activity,
betaxanthins. cough treatment, nematicidal, liver and kidney function
etc. (c.f. Datta et al., 2017).
Pigments in the magenta coloured bracts of Bougainvillea
cvs. ‘Princess Marget Rose’, ‘Mrs. H.C. Buck’ and ‘Mary
MOLECULAR DIVERSITY
Palmer’ were resolved by paper electrophoresis into 5
bands, 3 pertaining to betacyanins and 2 to betaxanthim The genetic variation in plants is considered to be an
with different electrophoretic mobilities. The genes unlimited resource for introgression of foreign genes
responsible for the synthesis of betacyanins have been (Saghai Maroof et al., 1984). RAPD technique is suitable
suppressed in ‘Mary Palmer’, a mutant of ‘Mrs. H.C. for confirmation of parent-hybrid relationship. RAPD
Buck’. They are, however, partially activated by light and technique has been used in Bougainvillea to characterize
high temperature. This potentiality is almost lost in the germplasm, hybrids, sports, mutants at molecular level
‘Shubra’, in which no bands of betacyanins were (Chatterjee et al., 2007; Kumar et al., 2014; Hammad,
described (Kochhar and Ohri, 1977; Kochhar et al., 1979). 2009; Kumar et al., 2015). Molecular markers provide
Heuer et al. (1994) isolated and characterized Betacyanins more positive identification of varieties and cataloguing
from the bracts of Bougainvillea glabra by a combination varieties according to molecular difference. Chatterjee et
of spectroscopic techniques as gomphrenin I (betanidin al. (2007) made extensive studies on RAPD analysis of

Volume 10, Number 2, May-August, 2021 69

 S.K. Datta

92 cultivars and the technique was found to be very helpful Development of New Varieties
for assessment of genetic diversity of large germplasm
Creation of genetic variability is prerequisite for
collections and identification of cultivars, documentation,
development of new variety. CSIR-NBRI made
and to trace out the molecular affinity of origin of
concentrated efforts to improve/develop new varieties of
unknown group of Bougainvillea cultivars. In floriculture,
bougainvillea using a number of techniques. Understanding
such study is very important for better management,
of breeding system was very important before practical
identification of accessions and also in avoiding
breeding.
duplications or mislabeling of the genotypes. Such study
helps further improvement through selective breeding
programme. Srivastava et al. (2009) studied about 21 Breeding System in Bougainvilea
Bougainvillea cultivars (belonging to five species with one For genetic improvement, the first attempt was to
cultivar of the B. spectabilis species, five of B. glabra, understand the breeding system of bougainvillea. Heredity
four of B. peruviana, eight of B. x buttiana and three of and variation are controlled by the genetic system and
B. specto-glabra) through RAPD analysis to confirm the these basic information help to prepare most suitable
parentage of the hybrids. They conclude that to obtain breeding methodology for genetic improvement. Relevant
reliable identification, tracing genetic relationships and cytogenetic characteristic were studied to understand the
characterization of the Bougainvillea germplasm, the mechanisms for domestication of bougainvillea. The first
molecular approach based on RAPD profile is a powerful step was to develop suitable genotype with different
technique. Genetic diversities and phylogenetic objectives through selection pressures. Selection pressure
relationships of 48 Bougainvillea germplasm belonging to along with genetic-evolutionary influence helped to develop
B. glabra and B. spectabilis, B. peruviana, B. × buttiana new variations in bougainvillea. There was necessary to
and B. × spectoglabra were studied based on RAPD and understand the breakdown in the original mating system
ISSR markers (Yuanjiang et al. 2013). Tang et al. (2014) of elemental species of bougainvillea with changing
analysed genetic diversity and relationship among 48 environmental conditions, from wild to the cultivated form.
Bougainvillea cultivars using sequence-related amplified
Khoshoo (1981) proposed such excellent knowledge in
polymorphism markers. Genetic diversity, genetic
breeding system of ornamentals in general and bougainvillea
relatedness and molecular characterization of fifty known/
in particular. Studied clearly indicate that most of the
unknown origin bougainvillea cultivars belonging to four
present day cultivars of bougainvillea have developed from
major species of bougainvillea (B. glabra, B. spectabilis,
four basal species namely B. glabra Choisy, B. peruviana
B. peruviana and Bougainvillea × buttiana) were
Humboldt and Bonpland, B. spectabilis Willdenow and B.
determined using SSR markers (Kumar et al., 2014).
x buttiana Holttum and Standley. The mechanisms
Hammad (2009) determined the genetic relationships
involved were natural and selective hybridization,
among the commonly grown cultivars in Egypt using
chromosomal manipulations and spontaneous and induced
RAPD isozymes pattern. Chemotaxonomy was studied
mutations. At the early stage a good amount of variation
to determine the inter-relationship of Bougainvillea with
was developed due to natural interspecific hybridization
other genera and species (Selvam and Rajasekaran 1993).
among the three species of bougainvillea. Further
Large genetic diversity among different germplasm
development of novel colours and interesting chimeric
resources was observed through determination of esterase
designs and double bracted cultivars through bud-sports
and peroxidase isozyme using polyacrylamide gel
from these hybrids.
electrophoresis (Chen et al., 2012; Chen et al., 2012;
Datta et al., 2017). Wu and Tang (2012) established Pollen and/or seed sterility in bougainvillea is a major
SRAP-PCR reaction system for Bougainvillea and utilized handicap for an extensive breeding programme. There is
the data for genetic analysis, map construction, gene restriction in selection of male and female parents for
localization and identification of germplasm. breeding. CSIR-NBRI did excellent work to use

70 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

Chromosome manipulation as a good tool for development vegetative means. Spontaneous bud variations have been
of new varieties. Fertility could be restored by colchicines- responsible for three types of changes in cultivated
induced polyploidy which helped development of very bougainvilleas viz. change in bract colour, development
desirable, colourful and floriferous, often bicoloured, of imperfect floral tubes and leaf variegation (Zadoo et
hybrid bougainvillea (triploid, tetraploid and aneuploid). al., 1975d).
Induced tetraploidy revealed a new outlook in bougainvillea
breeding. Triploid varieties were developed by crossing Classical Breeding
tetraploid with diploid. Bougainvillea cvs. ‘Shubhra’, ‘Mrs.
Plant breeding is one of the best methods of developing
McClean’, ‘Mary Palmer’ and ‘President Rosevelt’ are
new combinations. Because it helps in collection,
diploid and sterile. Tetraploidy was induced in these
incorporation and rearrangements of genetic diversity
varieties. Tetraploids are self incompatible which after
through selection. For undertaking a meaningful
crossing with other fertile forms produce seeds.
improvement programme, it is most imperative that the
Development of very promising cultivars at NBRI viz.
basic genetic information is obtained through the study
‘Begum Sikander’, ‘Wajid Ali Shah’, ‘Mary Palmer
of breeding systems, experimental hybridization and
Special’ and ‘Chitra’ by hybridizing ‘Dr. B.P. Pal’
synthesis of aneuploids and amphiploids, involving both
(tetraploid of ‘Shubhra’) and ‘Tetra Mrs. McClean’
the cultivated and elemental species from the wild. Such
(tetraploid of ‘Mrs. McClean’) with other sterile, diploid
studies, carried out at NBRI, have facilitated charting out
cultivars are worth mentioning (Khoshoo and Zadoo, 1969;
genetic evolutionary race histories of several important
Zadoo et al.,1975c; Khoshoo, 1990; Ohri, 2013).
crops including Bougainvillea. The information generated
Generally Bougainvillea cultivars are sexually sterile and by such studies have helped in the circumscription of
are multiplied by vegetative propagation. Some cultivars ‘gene pools’ and their utilization in the creation of new
produce seeds occasionally. Planned hybridization and novel cultivars of commercial importance keeping in
between the cultivars producing seeds and those having view the direction of market trend. Voluminous work have
fertile pollen is in practice for evolving new cultivars. been done on bougainvillea at CSIR-NBRI on breeding
Bougainvillea has comparatively simpler genetic system for development of new varieties through selective (known
as all the three elemental species are diploid and the hybrids parentage) breeding (Khoshoo, 1968, 1971a, b, 1981;
between them are either completely sterile or semi sterile. Banerji and Datta, 1993; Sharma, 1986; Sharma and
This interspecific hybridization has produced intermediate Srivastava, 1988, 1989; Zadoo and Khoshoo, 1968;
morphological and phenological characters and entirely Khoshoo and Zadoo, 1969; Zadoo et al., 1975b; Khoshoo,
novel bract colours in the three hybrid groups. The fertility 1990).
in these hybrid groups could be restored by induced
polyploidy and this expanded germplasm was used to Induced Mutagenesis for Development of New
produce tetraploid and triploid hybrids with highly positive Varieties
horticultural traits. Any novelty developed through any
CSIR-NBRI is one of the pioneer institutions where
technique could be established by vegetative multiplication
commendable work has been done on induced
(Ohri, 2013; Zadoo et al., 1975b).
mutagenesis. Appreciable information have been
accumulated on different aspects related to induced
Spontaneous Mutation (Bud sport)
mutations. Different mutagens like Gamma radiation, Ethyl
Spontaneous bud sports occur commonly in the cultivated Methane Sulphonate (EMS), Methyl Methane sulphonate
ornamental plants and account for considerable variation. (MMS) etc. have been used. Different single and double
Bud sports have been noted from time to time by keen bracted varieties were differentially sensitive to mutagens.
gardeners, horticulturists, researchers etc. in their Sensitivity of different bougainvillea genotypes to gamma
germplasm collections which are being maintained by radiation varied from 250 to 1250 rads and studies end

Volume 10, Number 2, May-August, 2021 71

 S.K. Datta

result concluded that radiosensitivity in Bougainvillea is a 2000). Sodium azide induced leaf colour and leaf shape
genotype dependent mechanism (Datta, 2009; Hong and mutations. Radiosensitivity studies on Bougainvillea have
Shaode, 1990). been reported (Chen et al., 2012; Swaroop et al., 2015).

It was curious to mention that mutation breeding

Detection, Isolation and Management of Chimera
experiments carried out at CSIR-NBRI resulted mutations
mostly in leaf variegations. No new bract colour mutations All the mutation/s (chlorophyll variegations in leaves or
could be induced and isolated except in cultivar ‘Palekar’. bract colour) in bougainvillea appeared as chimera.
The original young bract colour of ‘Palekar’ is Current Isolation of mutant tissues from chimeric branch is the
Red (821/3) changing to Salferino purple (26/1) on most important operation in mutation breeding. Normally
maturity. The mutant bract colour at young stage is Orange mutations appear mericlinal chimera. Mericlinal chimeras
(12/1) which on maturity becomes Azalia Pink (523/1). are unstable and there is always change in frequency of
Leaves in original cultivar are obtuse and flat while in normal and chlorophyll variegated branch. Periclinal
mutant these are elliptic and incurved. Present mutant of chimeras are stable and most commercial chlorophyll
‘Palekar’ is perhaps the first established gamma ray variegations are periclinal chimeras. Conventional
induced bract colour mutation in bougainvillea (Sharma technique has been standardized in bougainvillea for
et al., 2002). management of such chimera and development of new
varieties. In chimeric chlorophyll variegated branch, the
Mutation breeding of multibracted bougainvillea need
growth of the axillary bud associated with the variegated
special mention. Only four cultivars viz. ‘Cherry
leaves are arrested by the influence of the apical dominance
Blossom’, ‘Los Banos Beauty’, ‘Mahara’ and ‘Rosevilles
of the main apex of the mericlinal shoot. Simple technique
Delight’ comprise the original double bracted group.
has been standardized to convert mericlinal chimera to
Mutation breeding is the only method for improvement
periclinal chimera. It helps to isolate mericlinal branch to
of double bracted Bougainvillea due to absence of flower.
pure chlorophyll variegated branch (periclinal). By this
A total of four chlorophyll variegated mutants have been
technique new chlorophyll variegated mutants have been
developed - ‘Pallavi’, ‘Mahara variegata’, ‘Los Banos
isolated in pure form and commercialized in bougainvillea
Variegata’ and ‘Los Banos Variegata Jayanthi’. As far as
and other crop (Banerji and Datta, 1987; Datta, 1992).
brcat colour is concerned still no new bract colour could
be induced. It was interesting to note that although there
Origin of Cultivars in India
was formation of no new bract colour but formation of
chimera of existing bract colour was recorded. Irradiated Bougainvillea varieties available in India have been
population of ‘Roseville’s Delight’ produced chimeric bract mentioned above. An attempt has been made to trace out
of ‘Mahara’, ‘Cherry Blossom’ and ‘Los Banos Beauty’ the mode of origin of different varieties as per available
type; ‘Cherry Blossom’ produced ‘Mahara’ and ‘Los information. The exact origin of all the varieties are not
Banos Beauty’ type and ‘Los Banos Beauty’ produced available. Breeders, sometimes, do not disclose the origin
only ‘Mahara’ type chimera. But surprisingly no change of varieties. Information available is not complete in most
in bract colour in irradiated population of ‘Mahara’ was cases. New varieties of bougainvillea have been developed
recorded (Banerji and Datta, 1987). Abraham and Desai in India at different research institutions, nurseries,
(1977) developed four gamma ray induced mutants - botanical societies and also by individual efforts. Details
‘Jaya’, ‘Jayalaxmi variegated’, ‘Lady Hudson of Ceylon of origin and parentage of only few varieties are known.
variegata’ and ‘Silvertop’. New flower colour mutations As per available reports of different workers, varieties
have been reported in Bougainvillea spectabilis treating have been categorized as Hybrid Seedling, Seedling
with 60Co  rays (Deng and Liu 1990). EMS treatment Selection, Bud Sports, Mutant Variety, Chromosome
produced chlorophyll variegated leaf mutation in double manipulation, Individual Name etc. Names of all these
bracted variety ‘Los Banos Variegata’ (Jayanthi et al., new varieties and the total number (percentage) have been

72 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

Table 1. Name of cultivars and their probable origin

Hybrid Seedling [26.89%]
‘Aida’ (B. spectabilis, Lancaster, Calcutta), ‘Amarnath’ (Lancaster, Calcutta), ‘Amarault’ (Lancaster, Calcutta), ‘Autumn’ (Lancaster, Calcutta),
‘B.A. Rama Rao’ (Madras), ‘Beauty’ (Lancaster, Calcutta), ‘Begum Ali Yawar Jung’ (B. buttiana, Habib-ur-Rehman, Aligarh), ‘Begum Sikander’
(‘Dr. B.P. Pal’ x ‘Jannifer Fernie’, Zadoo and Khoshoo, NBRI, Lucknow), ‘Bonfire’ (Lancaster, Calcutta), ‘Brilliant Variegata’ (B. buttiana
cv. Brilliant, Lal Baugh Botanic Gardens), ‘Cascade’ (Agri-Horticultural Society, Madras), ‘Chandrabieri’ (B. buttiana, M/s. Chandra Nursery,
Sikkim), ‘Cherry Ripe’ (Lancaster, Calcutta), ‘Chitra’ (Tetra Mrs. McClean – B. buttiana x ‘Dr. B.P. Pal’ – B. peruviana, Khoshoo et al.,
NBRI), ‘Chitravati’ (B. spectabilis x ‘Red Glory’, IIHR, Bangalore), ‘Conquest’ (Lancaster, Calcutta), ‘Crimson King’ (M/s. Soundary Nursery,
Madras), ‘Crimson Lake’ (B. buttiana, Lancaster, Calcutta), ‘Dogstar’ (B. spectabilis, Lancaster, Calcutta), ‘Dr. H.B. Singh’ (B. glabra x
‘Formosa’, IIHR), ‘Dr. R.R. Pal’ (Thomasii – B. spectabilis x ‘Louise Wathen’ – B. buttiana, Dr. B.P. Pal, Delhi), ‘Durga’s Delight’ (M/s.
Chandra Nursery, Sikkim), ‘Eclipse’ (B. spectabilis, Lancaster, Calcutta), ‘Exquisite’ (Lancaster, Calcutta), ‘Flame’ (B. spectabillis, Soundarya
Nuesery, Madras), ‘Intermedia’ (Lancaster, Calcutta), ‘Lady Hope’ (B. peruviana, B.A.Rama Rao, Madras), ‘Lady Mountbatten’ (B. spectabilis,
Imperial Nursery, Calcutta & M/s Soundarya Nursery, Madras), ‘Lord Willingdon’ (‘Maharaja of Mysore’, Lancaster, Calcutta), ‘Maharaja of
Mysore’ (‘Lateritia’ – B. spectabili x B. glabra, B.S. Nirody), ‘Manohar Chandra’ (M/S. Chandra Nursery), ‘Mary Palmer Special’ (‘Dr B.P.
Pal’ – B. peruviana x ‘Princess Margaret Rose’ – B. peruviana, Zadoo and Khoshoo, NBRI), ‘Mrs. Fraser’ (B. spectabilis, B.S. Nirody,
Bangalore, 1932), ‘Mrs. Chico’ (Lancaster, Delhi), ‘Nawab Ali Yawar Jung’ (‘Habib-ur-Rehman’, Aligarh), ‘New Red’ (Agri-Horticultural
Society of India, Calcutta), ‘Nirmal Chandra’ (M/s Chandra Nursery, Sikkim), ‘Pink Beauty’ (B. glabra x ‘Lady Mountbatten’, M/s Soundary
Nursery), ‘Pradhan’s Profusion’ (M/s Chandra Nursery, Sikkim), ‘Purple Gem’ (‘Mrs.H.C. Buck’ – B. peruviana x ‘Refulgens’ – B. spectabilis,
M/s Soundary Nursery), ‘Spitfire’ (perhaps B. spectabilis, Lancaster, Calcutta), ‘Spring Festival’ (‘Thomasii’ – B. spectabilis x ‘Louise Wathen’
– B. buttiana, Pal, New Delhi), ‘Srimati Biezli’ (M/s Chandra Nursery), ‘Summer Time’ (‘Thomasii’ – B. spectabilis x ‘Louise Wathen’ – B.
buttiana), ‘Versicolour’ (B. buttiana, M/s Soundarya Nursery), ‘Vesuvius’ (Lancaster, Calcutta), ‘Vijaya’ (B. peruviana, M/s Soundarya Nursery),
‘Wajid Ali Shah’ (‘Dr. B.P. Pal’ – B. peruviana x ‘Mrs. Chico’ –B. spectabilis, Zadoo and Khoshoo, NBRI), ‘Win Some’ (Lancaster, Calcutta),
‘Purple Rose’ (Lancaster, Calcutta), ‘Purple Wonder’ (‘Formosa’ – B. glabra x ‘Trinidad’ – B. glabra, IIHR), ‘Ranee’ (M/s Soundary Nursery,
Madras), ‘Rosa Multiflora’ (‘Mrs. Fraser’ – B. spectabilis x ‘Louise Wathen’ – B. buttiana, M/s Soundary Nurserys), ‘Rose Queen’ (B. glabra
x ‘Lady Mountbatten’ – B. spectabilis, M/s Soundary Nursery), ‘Star’ (Lancaster, Calcutta), ‘Sweet Heart’ (M/s Soundarya Nursery, madras),
‘Udai Chandra’ (B. glabra, M/s Chandra Nursery, Sikkim), ‘Wajid Ali Shah’ (‘Dr. B.P. Pal’ X ‘Mrs. Chico’. Triploid., Zadoo and Khoshoo,
NBRI).
Bud Sports [31.60%]
‘Abhimanya’ (‘Arjuna’, Banerji, NBRI), ‘Alic Lancaster’ (‘Scarlet Queen’, B. buttiana, Lancaster, Delhi), ‘Anindita’ (Refulgens, S. Ghosh,
Kandi, West Bengal), ‘Archana’ (Roseville’s Delight, B. buttiana, NBRI, Srivastava), ‘Bhabha’ (Louise Wathen, B. buttiana, Lalbaugh Botanic
Garden), ‘Daya’ (Partha, B. peruviana, Palekar & Co., Bombay), ‘Enid Lancaster’ (‘Louise Wathen’, B. buttiana, Lancaster, Delhi), ‘Fantasy’
(‘Peincess Margaret Rose’, B. peruviana, Rama Rao, Madras), ‘Gangamma’ (‘Lady mary Baring’, B. buttiana, Munivenkatappa, Bangalore),
‘Gangaswamy’ (‘Lady Mary Baring’, B. buttiana, Lalbaugh Botanic Gardens), ‘Gem’ (B.S. Nirody, Indian Institute of Science, Bangalore),
‘Glabra variegatea’ (‘B. glabra’, Lal Baugh Botanic Gardens, Bangalore from Kenya), ‘Godrej Cherry Blossom’ (‘Cherry Blossom’), ‘Hawaiian
Beauty’ (‘Hawaiian White’, Sharma), ‘Jane Stansfeld’ (Refulgens, B. spectabilis, Rama Rao, Madras Club, Madras), ‘Jawaharlal Nehru’ (B.
spectabilis, IIHR), ‘Jayalakshmi Variegata’ (‘Jayalakshmi’, B. buttiana, Abraham and Desai, BARC), ‘Kuvempu’ (‘Rao’, B. buttiana, Lalbaugh
Botanic Gardens), ‘Lady Hudson of Ceylon Variegata’ (‘Lady Hudson of Ceylon’, Abraham and Desai, BARC), ‘Laxminarayana’ (B. spectabilis,
Lalbaugh Botanic Gardens, Bangalore), ‘L.N. Birla’ (‘Maharaja of Mysore’, Agri-Horticultural Society of India, Calcutta), ‘Louise Wathen’
(‘Mrs. Butt’ or ‘Scarlet Queen’, Nirody, Agri-Horticultural Society, Madras), ‘Louise Wathen Mediopicta’ (‘Louise Wathen’, ‘B. buttiana’,
Royal Agri-Horticultural Society of India, Calcutta), ‘Louise Wathen Variegata’ (‘Louise Wathen’, B. buttiana, Royal Agri-Horticultural
Society of India), ‘Magenta Queen’ (‘Mrs. Butt’, Rama Rao, Madras), ‘Mahatma Gandhi Variegated’ (‘Mahatma Gandhi’, Pratap Nursery &
Seed Store, Dehradun), ‘Manohar Chandra Variegata’ (‘Manohar Chandra’, Sharma and Bisaria, NBRI), ‘Marietta’ (‘Mahara’, Mrs. Marietta,
Rizal of Philippines, reported by Dr. J.V. Pancho), ‘Marigowda’ (‘Gladys Hepburn’, Lalbaugh Botanic Gardens), ‘Mary Palmer’ (‘Mrs. H.C.
Buck’, B. peruviana, Lancaster, Calcutta), ‘Mary Baring’ (‘Mrs. McClean’), ‘Mataji Agnihotri’ (‘Mary Palmer’, B. peruviana, M/s K.S.
Gopalaswamiengar & Son), ‘Meera Sport’ (‘Meera’, Imperial Nursery, Calcutta), ‘Midget’ (Lancaster, Calcutta), ‘Mrs. Butt Magenta’ (‘Mrs.
Butt, Lancaster, Lucknow), ‘Mrs. Butt Scarlet’ (‘Mrs. Butt’, Lancaster, Lucknow), ‘Mrs. G.S. Randhwa’ (‘Killie Campbell’, B. buttiana,
Randhawa, Delhi), ‘Mrs. McClean’ (‘Mrs. Butt’), ‘Mrs. McClean Nirmal’ (‘Mrs. McClean’, B. buttiana, G.S. Srivastava, NBRI),
‘Munivenkatappa’ (Mary Palmer’, B. peruviana, Y. Munivenkatappa, Nurseryman, Bangalore), ‘Nirmal’ (‘Mrs. McClean’, NBRI,
Srivastava),‘Odisee’ (‘Mary Palmer’, B. peruviana, Das, Orissa University of Agric. & Tech.), ‘Parthasarathy’ (‘Partha’, B. peruviana,
Sharma, NBRI), ‘Partha Variegata’ (‘Partha’), ‘Preeti’ (‘Partha’, Palekar & C0., Bombay), ‘Purple Prince’ (‘Mrs. Butt’, Rama Rao, Madras),
‘Purple Queen’ (‘Mrs. Butt’, Rama Rao, Madras), ‘Rao’ (‘Mrs. Butt, Lalbaugh Botanic Gardens, Bangalore), ‘Refulgens variegata’ (‘Refulgen’,
Gopala Swamy Iyengar sons, Bangalore), ‘Red Glory Improved’ (‘Red Glory’, M/s Gopalaswamiengar & Son), ‘Scarlet Glory’ (‘Padmi’, B.
buttiana, /s Gopalaswamiengar & Son), ‘Scarlet Queen Variegated’ (‘Scarlet Queen’, Lancaster, Calcutta), ‘Sharma’ (‘Mrs. Fraser’, Lalbaugh

Volume 10, Number 2, May-August, 2021 73

 S.K. Datta

Table 1 contd...

Botanic Gardens), ‘Shweta’ (‘Trinidad’, Sharma, NBRI), ‘Shubhra’ (‘Mary Palmer’, Sharma, NBRI), ‘Soundarya’ (‘Mrs. H.C. Buck’, M/s
Soundary Nursery), ‘Spectabilis Variegated’ (B. spectabilis, Lancaster, Lucknow), ‘Surekha’ (‘Scarlet Queen’, Sharma, NBRI), ‘Thimma’ (‘Mary
Palmer’, B. peruviana, Lalbough Botanic Gardens), ‘Vellayani’ (‘Mrs. Butt’, Lalbaugh Botanic Gardens), ‘Vishakha’ (‘Mrs. H.C. Buck’, B.
peruviana, IIHR), ‘Zakerina’ (‘Maharaja of Mysore’, B. specto-glabra, Lalbaugh Botanic Gardens), ‘Zakir Hussain’ (‘Maharaja of Mysore’,
Lalbaugh Botanic Gardens). ‘Purple King’ (‘Mrs. Butt’, Rama Rao, Madras), ‘Shweta’ (‘Trinidad’, B. glabra, Sharma, NBRI), ‘Suvarna’
(‘Lady Hudson of Ceylon’, B. peruviana, Desai and Abraham, BARC), ‘Yellow Queen’ (‘Mrs. McClean’, B. buttiana, Marigowda, Lalbaugh
Botanic Gardens).
Seedling Selection [20.75%]
‘Baby Margaret Rose’ (‘Princess Margaret Rose’, B. peruvialla, B. Ranga Reddy, Madras), ‘Brenda Ohare’ (‘Gopal’, B. glabra, Mrs. Verna
Nagpal), ‘Buddhadas’ (‘Snow Queen’ B. glabra, Yediur Botanic Gardens, Bangalore, M. Buddhadas), ‘Celia Braganza’ (Verna Nagpal), ‘Deep
Cherry’ (B. spectabilis, Agri-Horticultural Society, Calcutta), ‘Dream’ (B. glabra, Swaminathan, Soundary Nursery, Madras), ‘Eisha’ (‘Formosa’
– B. glabra, Mrs. Verna Nagpal, Bombay), ‘Ethiraj’ (B. glabra, Agri-Horticultural Society, Madras), ‘Gagarin’ (‘Trinidad’ – B. glabra, Lalbaugh
Botanic Gardens), ‘Gopal’ (‘Formosa’, B. glabra, M/s K.S. Gopalaswamiengar Son), ‘Happiness’ (‘Lateritia’, B. spectabilis, M/s
Gopalaswamiengar & Son), ‘Jayalakshmi’ (‘Meera’, B. buttiana, M/s Gopalaswamiengar & Son), ‘Jennifer Nagpal’ (‘Gagarin’, B. glabra,
Mrs. Verna Nagpal, introduced by M/s Gopalaswamiengar & Son), ‘Jubilee’ (B. spectabilis, Lancaster, Calcutta), ‘Kalyani’ (‘Gopal’, B. glabra,
Kasturi Rangan, M/s Gopalaswamiengar & Son), ‘Krumbiegal’ (‘Mahatma Gandhi’, B. peruviana, M/s Gopalaswamiengar & Son), ‘Lalbagh’
(‘Lateritia’, B. spectabilis, Lalbaugh Botanic Gardens), ‘Lilacina’ (B. glabra, M/s Soundarya Nursery), ‘Lilac Queen’ (‘Formosa’, B. glabra,
M/s Gopalaswamiengar & Son), ‘Lilac Perfection’ (B. glabra, M/s Gopalaswamiengar & Son), ‘Lilac Puff’ (B. glabra, M/s Gopalaswamiengar
& Son), ‘Louise Wathen Intermediate’ (B. buttiana, listed by Imperial Nursery, Calcutta), ‘Mahatma Gandhi’ (‘Princess Margaret Rose’, B.
peruviana, M/s K.S. Gopalaswamiengar & Son), ‘Meera’ (B. glabra, Rangachari, Madras), ‘Mrs. H.C. Buck’ (‘Princes Margaret Rose’, B.
peruviana, Swaminathan, M/s Soundary Nursery), ‘Mrs. Lancaster’ (‘Maharaja of Mysore’, B. specto-glabra, Lancaster, Calcutta), ‘Mrs.
R.B. Carrick’ (B. spectabilis, Rama Rao, Madras), ‘Mundanna’ (‘Snow Queen’ - B. glabra x ‘Trinidad’ – B. glabra, M/s Mudanna & Sons),
‘Orange Sceptre’ (‘Brasiliensis’, B. spectabilis, M/s K.S. Gopalaswamiengar & Son), ‘Padmi’ (‘Mrs. Butt’, M/s Gopalaswamiengar & Son),
‘Partha’ (‘Princess Margaret Rose’, B. peruviana, M/s Gopalaswamiengar & Son), ‘Raman’ (B. glabra, Lalbaugh Botanic Gardens), ‘Red
Glory’ (‘Mrs. Fraser’, B. spectabilis, M/s Gopalaswamiengar & Son), ‘Sachindananda’ (‘Snow Queen’, B. glabra, M. Sachidananda, Yediur
Botanic Gardens), ‘Sholay’ (‘Red Glory’, IIHR), ‘Simson Anselm’ (‘Gem’, Verna Nagpal), ‘Snow Queen’ (B. glabra, introduced in India by
Dr. Homi Bhabha, Bombay), ‘Sonnet’ (B. buttiana, Pal, New Delhi), ‘Srinivasa’ (‘Jayalakshmi’, B. buttiana, M/s Gopalaswamiengar & Son),
‘Stanza’ (B. glabra, Pal, New Delhi), ‘Trinidad’ (‘Snow Queen’, B. glabra, Marigowda, Lalbaugh Botanic Gardens), ‘Usha’ (‘Lady Hope’, B.
peruviana, IIHR), ‘Verna Nagpal’ (‘Gagarin’, B. glabra, Nagpal, Bombay), ‘Villayani Variegata’ (B. buttiana, Vittal, Madras Municipal
Corporation Garden).
Mutant variety [6.60%]
‘Arjuna’ (‘Partha’, NBRI, Gupta and Shukla), ‘Aruna’ (Leaves light green, curled inwards), ‘Jaya’ (‘Jayalakshmi’, Abraham and Desai, BARC),
‘Jayalaxmi variegated’ (‘Jayalaxmi’, Abraham and Desai, BARC), ‘Lady Hudson of Ceylon Variegata’ (‘Lady Hudson of Ceylon’, Abraham
and Desai, BARC), ‘Los Banos Variegata silver margin’ (‘Los Banos Beauty’, NBRI, Lucknow), ‘Los Banos Variegata ‘Jayanti’ (‘Los Banos
Beauty’, NBRI, Lucknow, Jayanti et al.), ‘Mahara Variegata’ (‘Mahara’, NBRI, Datta and Banerji), ‘Mahara Variegata abnormal leaves’
(‘Mahara’, NBRI, Lucknow), ‘Dr. P.V. Sane’ (NBRI), ‘Silver Top’ (‘Versicolour’, B. buttiana, Abraham and Desai, BARC), ‘Pallavi’ (‘Roseville’s
Delight’, NBRI, Lucknow, Banerji et al.), ‘Pixie Variegata’ (NBRI), ‘Poultoni Variegata’ (‘Poultoni’, B. butiana, Abraham and Desai, BARC,
Bombay).
Individual Name [13.20%]
‘Crimson Rose’ (Lancaster, Calcutta), ‘Dwarf Gem’ (Lancaster, Calcutta), ‘Dorothy Jivarajadasa’ (Madras), ‘Glorious’ (M/s Soundarya Nursery,
Madras), ‘Gokul’ (M/s Soundary Nursery), ‘Gulaby’ (M/s Soundarya Nursery), ‘Hiawatha’ (Lancaster, Lucknow), ‘Jasper Rose’ (Agri-Horticultural
Society, Madras), ‘Lilac Beauty’ (M/s Gopalaswamiengar & Son, Bangalore), ‘Mauve Queen’ (M/s Soundary Nursery), ‘Mrs. Kay Malvenan’
(Rama Rao, Madras), ‘Mrs. Marie Buck’ (Rama Rao), ‘Orange Glory’ (Lancaster, Calcutta), ‘Pearl’ (M/s A. Boseck & Co., Calcutta), ‘Perfection’
(M/s Soundary Nursery), ‘Pradhan’s Pink’ (M/s Chandra Nursery, Sikkim), ‘President Rosevelt’ (M/s Soundary Nursery),’Princess Margaret
Rose’ (Agri-Horticultural Society, Madras), ‘Profusion’ (Lancaster, Calcutta), ‘Purple Mrs. Butt’ (M/s Gopalaswaniengar & Son), ‘Purple
Star’ (M/s Soundary Nursery), ‘Ratna’ (M/s Chandra Nursery), ‘Rosea Fuchsea’ (Agri-Horticultural Society, Madras), ‘Sensation’ (M/s Soundary
Nursery), ‘Star Mauve’ (Bose, Royal Agri-Horticultural Society of India, Calcutta), ‘Dr. B.P.Pal’, (New Delhi), ‘Superb’ (M/s Soundarya
Nursery), ‘Tyrian Rose’ (Agri-Horticultural Society, Madras), ‘Vijayalakshmi’ (M/s Gopalaswamiengar & Son), ‘Vithal Variegata’ (Madras
Municipal Corp. Garden).
Tetraploid [0.94%]
‘Dr. B.P. Pal’ (tetraploid of Shubhra – B. peruviana, Zadoo and Khoshoo, NBRI), ‘Tetra Mrs. McClean’ (tetraploid of ‘Mrs. McClean’, B.
buttiana, Zadoo and Khoshoo, NBRI).

74 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

shown in Table 1. Indian Institute of Horticultural Different bougainvillea genotypes are supposed to be very
Research, Bangarore developed variety ‘Dr. H.B. Singh’ sensitive to bud sport. Therefore, germplasm collection
got patent in Australia in the name of ‘Krishna’ (Dr. H.B. should be enriched and routine screening of germplasm
Singh : ‘Krishna’, Application No: 97/119 Accepted: 12 should be encouraged to detect minute changes as
Jan 1998) 4,28,56,87-95 (Sindhu, 2015; Sindhu and chimera. Now techniques are available for management
Swaroop, 2015; Singh et al., 1999; Nirody, 1932; Sharma, of such chimera into new form.
1963, 1964; Pancho, 1963, 1967; Marigowda, 1957;
In ornamentals, flower pigment is an important factor
Pancho and Elisco, 1959; Pancho and Capinpin, 1961;
which determines flower colour and its commercial value.
Srinivas, 1995). Following abbreviations have been used
Analysis of bract pigment of parental cultivars, sports,
– Hybrid Seedling (HS), Bud Sport (BS), Induced Mutant
hybrids and mutants of bougainvillea are very important
(IM) : (Table 1).
for future research. Pigment analysis will also help to
plan in vitro mutagenesis for developing new colour.
CONCLUSION
Conservation of genetic diversity of Bougainvillea is
Present day Bougainvillea represents collective diversity
important aspect. Because it has been realized that much
of different characters like other ornamentals
of the genetic variation available in the centres of diversity
(Chrysanthemum, Gladiolus, Rose etc.). Diversity
will soon become extinct due to several reasons if care is
developed in different parts of India through detection
not taken of it. Attention should be paid for both ex situ
and isolation of bud sports, hybridization and induced
and in situ conservation of bougainvillea germplasm at
mutation. All the basic scientific informations are available
different places.
on different aspects of Bougainvillea. But recently the
speed of development of new varieties has declined. No Bougainvillea is perhaps one ornamental plant where
further systematic research work is being carried out at remarkable changes have taken place in growth habit during
any research institute/university for further improvement. domestication through selection pressure and through
Proper efforts should be initiated by different organizations complex inter-specific crosses among elemental species
in this direction. and varieties and spontaneous mutation. But above all,
human selection pressure moulded bougainvillea for
The genetic diversity analysis will facilitate choosing the
multipurpose use. Climate change due to higher
appropriate breeding program to incorporate beneficial
temperature, changing rainfall and higher cabon dioxide
genes in desirable genotypes. We should initiate planned
level will affect future agriculture. Drought, saline and
experimental hybridization among the cultivated and
pollution tolerant are unique characters of bougainvillea.
elemental species. A good amount of variety specific
Work may be specially intensified for selection/
morpho-chemical characters and desirable genes have
development of more promising germplasm tolerant to
already been identified. Bougainvillea have many beneficial
such abiotic stress.
components for the consumer that can be created,
enhanced, or improved through breeding programme.
ACKNOWLEDGEMENT
Breeding objectives may be diversified as need and societal
benefit. The author thanks the Director CSIR-National Botanical
Research Institute, Lucknow, India for providing the
Appreciable breeding concepts have been reported by
working facilities.
CSIR-NBRI and different breeders and amateur growers.
But unfortunately such wealth of information have not
REFERENCES
been utilized by successive breeders. Bougainvillea is
perhaps one ornamental where not much directive Abraham V and Desai BM, 1977. Radiation induced variegated
breeding work has been done in spite of having so much mutant in bougainvillea. Current Science, Vol. 46, No. 10,
pp. 351-352.
diversity.

Volume 10, Number 2, May-August, 2021 75

 S.K. Datta

Banerji BK and Data SK, 1993. Mrs Butt – Bud sports and its International Atomic Energy Agency, Vienna, Austria, pp.
near relatives. Scientific Horticulture (Ed. S.P. Singh), Scientific 300-302.
Publishers, Jodhpur. Datta SK, Jayanthi R and Janakiram T, 2017. Bougainvillea. New
Banerji BK and Datta SK, 1987. Gamma ray induced mutation in India Publishing Agency, New Delhi, India.
double bracted Bougainvillea cv. ‘Los Banos Beauty’. Journal Deng H and Liu S, 1990. The study of radiation induced breeding
of Nuclear Agriculture Biology, Vol. 16, No. 1, pp. 48-50. in Bougainvillea. Journal of South China Agricultural
Banerji SK and Banda GK, 1967. A natural triploid in University, Vol. 13, No. 1, pp. 89-93.
Bougainvillea. Indian Journal of Horticulture, Vol. 24, No. Erdtman G, 1952. Pollen morphology and plant taxonomy :
1, pp. 106-108. angiosperms. Alunqvist and Wiksell, Stockholm.
Begum J and Datta RM, 1971. Variations in chiasma frequency Gills WT, 1976. Bougainvilleas for cultivation (Nyctaginaceae).
in B. spectabillis wild and B. buttiana Linn. The glamorous Baileya, Vol. 20, No. 1, pp. 34-41.
garden ornamentals. Science and Culture, Vol. 37, No. 1, pp.
295. Hammad I, 2009. Genetic variation among Bougainvillea glabra
cultivars (Nyctaginaceae) detected by Rapd markers and
Chandel VS, Kumar R, Pandey P and Singh L, 2015. Foliage Isozymes patterns. Journal of Agriculture and Biological
variegation and flower colour varying shade beauty in Sciences, Vol. 5, No. 1, pp. 63-71.
bougainvillea varieties. Indian Bougainvillea Annual, Vol. 26,
Heimerl A, 1900. Monographie der Nyctaginaceen. I. Bougainvillea,
No. 2, pp. 21-22.
Phaeoptilum, Colignonia. – Denkschr. Kaiserl. Akad. Wiss.,
Chatterjee J, Mandal AKA, Chakrabarty D and Datta SK, 2007. Wien. Math.-Naturwiss. Kl, Vol. 70, No. 2, pp. 97–137.
Use of RAPD analysis to determine genetic diversity and
Heuer S, Richter S, Metzger JW, Wray V, Nimzt M and Strack
relationships among Bougainvillea cultivars at inter-specific
D, 1994. Betacyanins from bracts of Bougainvillea glabra.
levels. Horticulture, Environment, and Biotechnology, Vol. 48,
Phytochemistry, Vol. 37, No. 3, pp. 761-767.
No. 1, pp. 43-51.
Holttum RE, 1955a. The cultivated bougainvilleas II. Bougainvillea
Chen T, Liu W, Xie L and Lei J, 2012. Analysis on Genetic
x buttiana, its varieties and hybrids. MAHA Magazine, Vol.
Diversity of Bougainvillea spp. Germplasm Resources Based
12, pp. 2-11.
on Isozyme. Journal of Anhui Agricultural Sciences, Vol. 1,
No. 2, pp. 29-31. Holttum RE, 1955b. The cultivated bougainvilleas. The varieties
of B. glabra. Malayan Agriculture Magazine and Journal,
Chen T, Wang A, Liu Y and Liu W, 2012. The Study on Mutagenic
Vol. 12, No. 1, pp. 25-36.
Effect of Bougainvillea Induced by NaN3. Chinese
Agricultural Science Bulletin, pp. 25. Holttum RE, 1957. Bougainvillea ‘Mary Palmer’. Malayan Agri-
Horticultural Association, Vol. 14, No. 1, pp. 13.
Crane MB and Lawrence WJC, 1952. The Genetics of Garden
Plants, 4th ed. London. Hong D and Shaode L, 1990. The study of radiation induced
breeding in Bougainvillea. Journal of South China Agricultural
Datta SK and Banerji BK, 1995. Cytological studies in ornamental University, Vol. 11, No. 1, pp. 89-93.
plants. Advances in Horticulturevol. 12-Ornamental plants
Part I (eds. K.L. Chadha and S.K. Bhattacharjee), Malhotra Jana BK and Maithi RG, 1986. Graceful bougainvillea for multiple
Publishing House, New Delhi, pp. 267-334. use. Floriculture Today, Vol. 6, No. 2, pp. 1986.

Datta SK, 1992. Mutation studies on double bracted Bougainvillea Jayanthi R, 1999. Varietal characterization and improvement of
at National Botanical Research Institute (NBRI), Lucknow, Bougainvillea through induced mutation, Ph.D. thesis,
India. Mutation Breeding Newsletter, IAEA, Vienna, Vol. 12, Bhimrao Ambedkar University, Agra.
No. 39, pp. 8-9. Jayanthi R, 2000. Selecting bougainvillea cultivar for landscaping.
Indian Bougainvillea Annual, Vol. 15, No. 4, pp. 27-31.
Datta SK, 2004. Bougainvillea research at National Botanical
Research Institute, Lucknow. Journal of Ornamental Jayanti R, Datta SK and Verma JP, 2000. Chemical induced
Horticulture, Vol. 7, No. 1, pp. 1-4. mutation in double bracted bougainvillea cv. ‘Los Banos
Beauty’. Indian Bougainvillea Annual, Vol. 15, pp. 9-11.
Datta SK, 2009. Role of Classical Mutagenesis for Development
of New Ornamental Varieties. In : Induced Plant Mutations Khoshoo TN and Zadoo SN, 1969. New perspectives in
in the Genomics Era (Ed. Q.Y. Shu). Joint FAO/IAEA Bougainvillea breeding. Journal of Heredity, Vol. 60, No. 6,
Division of Nuclear techniques in Food and Agriculture, pp. 357-360.

76 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

Khoshoo TN, 1968. Genetic improvement of ornamentals in India Marigowda MH, 1961. Miscellaneous notes – Bougainvillea
: Prospects, experimental approaches and suggestions. Indian buttiana Holttum and Standley and its cultivars in Lalbaug.
Journal of Genetic, Vol. 28, No. 1, pp. 187-198. Lalbagh Journal of Mysore Horticultural Society, Vol. 6, No.
Khoshoo TN, 1971a. Taxonomy of cultivated plants : Principals, 2, pp. 23-26.
procedures and prospects. Science & Culture, Vol. 37, No. Nageshwar G and Radhakrishnaiah M, 1994. A note on the
1, pp. 313-315. chemosystematics of Boerhavia and Bougainvillea
Khoshoo TN, 1971b. Hortorium taxonomy. Indian Journal of (Nyctaginaceae). Fedder Repertorico, Vol. 105, No. (1-2), pp.
Genetics, Vol. 31, No. 2, pp. 305-315. 45-47.

Khoshoo TN, 1981. Breeding systems in relation to evolution in Nair PKK, 1961. Pollen grains of cultivated plants. II.
some ornamentals. Proceedings of the Indian National Science Bougainvillea Comm., Hibiscus Medik. and Euphorbia
Academy, Vol. 47, No. 2, pp. 279-301. pulcherima Willd. Journal of Indian Botanical Society, Vol.
40, No. 2, pp. 367-381.
Khoshoo TN, 1986. Perspectives in Bougainvillea breeding. News
Letter, Bougainvillea Society of India, Vol. 6, No. 2, pp. 7-10. Ninan T, Singh MP and Swaminathan MS, 1959. Meiotic
behaviour and pollen fertility in some varieties of
Khoshoo TN, 1990. Creativity in ornamentals. Indian Horticulture,
Bougainvillea. Journal of Indian Botanical Society, Vol. 38,
Vol. 34, No. 2, pp. 25-29.
No. 1, pp. 140-145.
Kobayashi DK, McConnell J and John Griffis J, 2007. Bougainvillea.
Nirody BS, 1932. Bougainvillea glabra var. Louise Wathen.
Ornamentals and Flowers, Vol. 2, No. 1, pp. 1-12.
Gardener’s Chronicle, Vol. 91, No. 2, pp. 11.
Kochhar VK and Ohri D, 1977. Bio-chemical analysis of bract
Ohri D and Khoshoo TN, 1982. Cytogenetics of cultivated
mutations in bougainvilleas I. ‘H.C. Buck’ family. Zeitschrift
Bougainvillea. Z. Pflanzenzucht, Vol. 88, No. 2, pp. 168-173.
fur pflanzenzüchtung, Vol. 79, No. 1, pp. 47-51.
Kochhar VK, Kochhar S and Ohri D, 1979. Bio-chemical analysis Ohri D and Zadoo SN, 1975. Cross-compatibility studies in some
of bract mutation in bougainvillea cv. Mrs. H.C. Buck. Bougainvillea cultivars. Incompatibility Newsletter, Vol. 6, No.
Current Science, Vol. 48, No. 4, pp. 163-164. 2, pp. 35-39.

Kumar G, Sindhu SS and Singh KP, 2007. Bougainvilleas : Varied Ohri D and Zadoo SN, 1979. Cytogenetics of cultivated
uses in landscaping. Indian Bougainvillea Annual, Vol. 20, Bougainvillea VIII. Cross compatibility relationship and
No. 1, pp. 17-18. origin of Bougainvillea ‘H.C. Buck’ family. Z. Pflanzenzucht,
Vol. 82, No. 1, pp. 182-186.
Kumar PP, Janakiram T, Bhat KV, Jain R, Prasad KV and Prabhu
KV, 2014. Molecular characterization and cultivar Ohri D and Zadoo SN, 1986. Cytogenetics of cultivated
identification in Bougainvillea spp. using SSR markers. The Bougainvillea IX. Precocious centromere division and origin
Indian Journal of Agricultural Sciences, Vol. 84, No. 9, pp. of polyploid taxa. Plant Breeding, Vol. 97, No. 2, pp. 227-
1124-1130. 231.

Kumar PP, Janakiram T, Bhat KV, Prasad KV and Jain R, 2015. Ohri D, 1995. Bougainvillea breeding. In: Advances in Horticulture,
Genetic divergence analysis of bougainvillea (Bougainvillea Vol. 12, No. 1, p. 56.
spp) cultivars using morphological markers. The Indian Journal
Ohri D, 2013. Cytogenetics of domestication and improvement
of Agricultural Sciences, Vol. 85, No. 5, pp. 661-615.
of garden Gladiolus and Bougainvillea. The Nucleus, Vol. 56,
Kumar R, 2001. Floral variability studies in bougainvillea. Indian No. 3, pp. 149-53. doi: 10.1007/s13237-013-0091-7.
Bougainvillea Annual, Vol. 16, No. 2, pp. 21–25.
Ornamental Plants, Part I. (Eds.). Chadha, K.L. and Bhattacharjee,
Kumar R, Prasad A, Chandel VS and Mishra LK, 2015. S.K. New Delhi, India : Malhotra Publishing House, 2015.
Bougainvillea : Its special qualities for choice planting. Indian
Pancho JV and Bardenas AE, 1959. Bougainvilleas in the
Bougainvillea Annual, Vol. 26, No. 1, pp. 18.
Philippines. Baileya, Vol. 7, No. 3, pp. 91-100.
López HA and Galetto L, 2002. Flower Structure and Reproductive
Pancho JV and Capinpin JM, 1961. Haploidy in bougainvillea.
Biology of Bougainvillea stipitate (Nyctaginaceae). Plant
Philippine Agricultural Scientist. Vol. 45, No. 2, pp. 88-94.
Biology, Vol. 4, No. 4, pp. 508-514.
Marigowda MH, 1957. Bougainvillea buttiana. Holttum and Pancho JV and Elisco AB, 1959. Bougainvillea in the Philippines
Standley and its cultivars in Lal Baugh Bangalore. Journal of Baileys. Journal of Horticultural Taxonomy, Vol. 7, No. 3,
Bombay Natural History Society, Vol. 54, No. 1, pp. 801-804. pp. 91-101.

Volume 10, Number 2, May-August, 2021 77

 S.K. Datta

Pancho JV, 1963. Notes on two outstanding new cultivars of Bougainvillea ‘Palekar’. Journal of Nuclear Agriculture
Bougainvillea in the Philippines. Lal Baugh Journal of Biology, Vol. 31, No. (3-4), pp. 206-208.
Mysore Horticulture Society, Vol. 8, No. 3, pp. 25-26.
Sharma, S.C., 1986. Horto-taxonomical studies on Bougainvilleas.
Pancho JV, 1967. Notes on double bracted bougainvillea in the Ph.D. Thesis, Kanpur University, Kanpur, India.
Philippines. Lal Baugh Journal of Mysore Horticultural
Sindhu SS and Swaroop K, 2015. Bougainvillea repository at IARI.
Society, Vol. 12, No. 12, pp. 30-32.
Indian Bougainvillea Annual, Vol. 26, No. 3, pp. 27-29.
Pramanick DD, Mondal M and Maiti GG, 2015. Pollen
Sindhu SS, 2001. Bougainvillea cultivars and their suitability in
morphological studies on some members of the family
landscaping. Bougainvillea News Letter, Vol. 16, No. 2, pp.
Nyctaginaceae in India. Asian Journal of Plant Science and
13-14.
Research, Vol. 5, No. 2, pp. 72-76.
Sindhu SS, 2015. Registration of new cultivars. Indian Bougainvillea
Saghai Maroof MA, Solinan KM, Jorgensen RA and Allard RW,
Annuals, Vol. 26, No. 3, p. 30.
1984. Ribosomal DNA spacer-length polymorphisms in
barley : Mendelian inheritance, chromosomal location, and Singh AP, 1996. Some popular varieties of bougainvillea.
population dynamics. Proceedings of the National Academy Bougainvillea Newsletter, Vol. 11, No. 1, pp. 8- 10.
of Sciences of the United States of America, Vol. 81, No. 2, Singh B, Panwar RS, Voleti SR, Sharma VK and Thakur S, 1999.
pp. 8014-8028. The New International Bougainvillea Check List. Indian
Selvam BA and Rajasekaran K, 1993. A numerical taxonomic Agricultural Research institute, New Delhi.
study of South Indian Nyctaginaceae. Proceedings of the Srinivas M and Bhat RN, 1982 Cytological studies in some
National Academy of Science, India, Section B. Biological variegated Bougainvillea. The Nucleus, Vol. 25, No. 3, pp.
Sciences, Vol. 63, No. 4, pp. 413-416. 130-135.
Sen SN and Sen NK, 1954. Chromosome number and abnormalities Srinivas M, 1995. Bougainvillea. In : Advances in Horticulture
in Bougainvillea. Did you mean: Bulletin of the Botanical Vol. 12 – Ornamental Plants (Eds. K.L. Chadha and S.K.
Society of Bengal, Vol. 8, No. 1, pp. 140-142. Bhattacharjee), Malhotra House, New Delhi, pp. 140-156.
Sharma AK and Bhattacharya NK, 1960. Cytological Srivastava R, Shukla S, Soni1 A and Anil Kumar A, 2009. RAPD-
investigations on Bougainvillea as an aid in interpreting the based genetic relationships in different Bougainvillea cultivars.
evolution and affinities of different species and varieties. The Crop Breeding and Applied Biotechnology, Vol. 9, No. 2, pp.
Nucleus, Vol. 3, No. 3, pp. 19-76. 154-163.
Sharma M D, 1963. Some bewitching Bougainvilleas from Lal Swaroop K, Jain R and Janakiram T, 2015. Effect of different
Bagh. Indian Horticulture, Vol. 7, No. 2, pp. 19-20. doses of gamma rays for induction of mutation in
Sharma MD, 1962. New bougainvillea introduced in 1961. Lal bougainvillea cv Mahatma Gandhi. Indian Journal of
Baugh Journal of Mysore Horticulture Society, Vol. 7, No. 1, Agricultural Sciences, Vol. 85, No. 9, pp. 1245–1247.
pp. 18-23. Swarup V and Singh B, 1964. Pollen morphology and leaf harirs
Sharma MD, 1964. Variegated leaf and multibracts – A new interest in classification of Bougainvillea. Indian Journal of
in Bougainvillea spectabilis. Delhi Garden Magazine, Vol. 3, Horticulture, Vol. 21, No. 2, pp. 155-164.
No. 1, pp. 11-13. Takahashi M and Skvarla JJ, 1991. Exine Pattern Formation by
Sharma SC and Srivastava S, 1988. Million dollar bougainvilleas. Plasma Membrane in Bougainvillea Spectabilis Willd.
News Letter, Bougainvillea Society of India, Vol. 7, No. 1, (Nyctaginaceae). American Journal of Botany, Vol. 78, No.
pp. 1-3. 8, pp. 1063-1069.

Sharma SC and Srivastava S., 1989. Bicoloured bougainvilleas. Tang Y, Wu X and Cao W, 2013. Genetic diversities and
News Letter, Bougainvillea Society of India, Vol. 8, No. 1, phylogenetic relationships of Bougainvillea germplasm
pp. 7-11. resources based on RAPD and ISSR markers. Acta Botanica
Boreali-Occidentalia Sinica, Vol. 33, No. 7, 1325-32.
Sharma SC, 1996. Bougainvilleas in India. Economic Botany
Information Service, National Botanical Research Institute, Tang Yuan-jiang Wu, Xiaoyan and Cao Wenjing, 2014. Genetic
Lucknow, India. Diversity and Relationship of Bougainvillea Germplasm
Resources Based on SRAP Markers. Journal of Tropical and
Sharma SC, Srivastava R, Datta SK and Roy R, 2002. Gamma
Subtropical Botany, Vol. 22, No. 2, pp. 147-154.
ray induced bract colour mutation in single bracted

78 LS - An International Journal of Life Sciences

 Genetic Diversity and Improvement of Bougainvillea

Wilson GB, 1947. Meiosis in Bougainvillea. Nature, London, Vol. fertility in sterile cultivars. Ehphytica, Vol. 24, No. 3, pp.
160, No. 4056, pp. 121. 517-24.
Wu X and Tang Y, 2012. Optimization of SRAP-PCR reaction Zadoo SN, Roy RP and Khoshoo TN, 1975d. Cytogenetics of
system and primer screening in Bougainvillea [J]. Journal of cultivated Bougainvillea III. Bud sports, Z. Pflanzenzuchtg,
Huazhong Normal University (Natural Sciences), Vol. 3, No. Vol. 74, No. 2, pp. 223-239.
3, p. 56
Zadoo SN, Roy RP and Khoshoo TN, 1975e. Cytogenetics of
Xu S, Huang Q, Shu Q, Chen C and Vick AB, 2009. Reproductive cultivated Bougainvillea IV. Meiotic system. La Cellule, Vol.
organography of Bougainvillea spectabilis Willd. Scientia 71, No. 4, pp. 311-322.
Horticulturae, Vol. 120, No. 3, pp. 399-405.
Zadoo SN, Roy RP and Khoshoo TN, 1975f. Cytogenetics of
Zadoo SN and Khoshoo TN, 1968. Cytological studies in cultivated Bougainvillea II. Pollination mechanism and
Bougainvillea. I – A case of interchange heterozygocity. breeding system. Proceeding of Indian Academy Science, Vol.
Genetica, Vol. 39, No. 2, pp. 353-360. 41, No. 2, pp. 498-502.
Zadoo SN and Khoshoo TN, 1975. Nature of incompatibility in Zadoo SN, Roy RP and Khoshoo TN, 1976. Cytogenetics of
cultivated Bougainvillea. Compatibility Newsletter, Vol. 5, No. cultivated Bougainvilleas VII. Origin and evolution of
2, pp. 73-75. ornamental taxa. Indian Journal of Horticulture, Vol. 33, No.
2, pp. 278-288.
Zadoo SN, Roy RP and Khoshoo TN, 1975a. Cytogenetics of
cultivated Bougainvilleas I. Morphological Variation. Zhou Q, Huang Kefu, Ding Yin-long and Guo Huizhu, 2011.
Proceedings of the Indian National Science Academy, Vol. 41, Investigation and Taxonomic Identification on Introduced
No. 2, pp. 121-132. Ornamental Varieties in Bougainvillea in China. Acta
Zadoo SN, Roy RP and Khoshoo TN, 1975b. Cytogenetics of Agriculturae Jiangxi, Vol. 23, No. 5, p. 25.
cultivated Bougainvillea VI. Hybridization. Z. Pflanzenzuchtg,
Vol. 75, No. 3, pp. 114-137. How to cite this article: Datta SK, 2021. Genetic Diversity
and Improvement of Bougainvillea. LS - An International Journal
Zadoo SN, Roy RP and Khoshoo TN, 1975c. Cytogenetics of of Life Sciences, Vol. 10, No. 2, pp. 61-79.
cultivated Bougainvillea V. Tetraploidy and restoration of

Volume 10, Number 2, May-August, 2021 79

View publication stats