Banana Resources

Banana Genetic Resources
S. Uma, M. S. Saraswathi, and P. Durai
Abstract Banana is a nature’s gift to the mankind with four times protein, twice
carbohydrates, three times phosphorus, five times vitamin A and iron, many times
potassium and twice other vitamins and minerals compared to apple. It is under
cultivation in more than 120 countries across the globe. It is referred as a food
equivalent across major African countries, Latin America including Caribbean and
Polynesian Islands. Therefore it has acquired the food-fruit status contributing
towards food and nutritional security. This chapter describes the botany and
taxonomy of banana, Musa genetic resources and their allied species available
across the globe, the evolutionary history of various ploidies and genomes. This also
provides elaborate practical guidelines for the successful collection of banana
genetic resources. This includes strategies for germplasm collection, establishment
of virus-free germplasm, characterization, conservation, documentation and their
potential utilization in banana improvement. This deals with the current status and
future research needs of various conservation strategies. This chapter recommends
the strategic collection of genetic resources and indexing prior to establishment with
special emphasis on application of advanced GIS tools in relation to ecological data
for effective conservation.
Keywords Banana · Genetic resources · Collection · Characterization ·

Conservation
1 History, Origin and Evolution
Banana (Musa spp.) with its versatile utilities is recognized as one of the earliest
crops domesticated by humans. As a wild seeded plant, banana must have been first
recognized for its nonedible purposes like fibre, roofing and navigation purposes.
S. Uma (*) · M. S. Saraswathi · P. Durai

ICAR – National Research Centre for Banana, Tiruchirapalli, Tamil Nadu, India
© Springer Nature Singapore Pte Ltd. 2019 321

P. E. Rajasekharan, V. R. Rao (eds.), Conservation and Utilization
of Horticultural Genetic Resources, https://doi.org/10.1007/978-981-13-3669-0_10
322 S. Uma et al.
Earliest documentary mention is in the Vedic period (approx. 1700 BC) where the
use of fruits like mangoes, goose berries and bananas has been mentioned in Rig-
Veda, one of the four volumes of Veda written by the Indian sages (Kulkarni 1973).
Next evidences are from Indian epics, ‘Mahabharata’ and ‘Ramayana’ which date
to approx. 1400 BC. Aranya Kanda and the forest trek of Valmiki’s ‘Ramayana’
mentioned about people of royal heritage draped in clothes woven in banana fibre.
Mention of banana fruits as contemporary diet in Ramayana indicated the domes-
ticated status of banana in the Indian subcontinent. Early epics of the Pali Buddhist
monks also mentioned about bananas ‘as big as an elephant’s tusk’ in 500–
600 BCE. This probably referred to plantains (Horn plantain?) from southern pen-
insula of Greater India (including Sri Lanka), where plantain cultivation is in vogue
today. Although it is an undisputed fact that banana and plantain originated in
Southeast Asia and the Pacific and it has been proved from chronology of their
spread and distribution, recent linguistics, anthropological and archaeological
studies have given indications on the possible migratory theory (Nayar 2010).
The genus Musa has been classified into four major sections, namely, Eumusa,
Rhodochlamys, Callimusa and Australimusa, but majority of the cultivated bananas
originated from Eumusa which is the biggest section in the genus and the most
geographically diverse, with species being found throughout Southeast Asia from
India to the Pacific Islands (Horry et al. 1997).
Though there are many species in the section Eumusa, only M. acuminata and
M. balbisiana are believed to have contributed to the evolution of present-day
cultivated bananas which are sterile and parthenocarpic (Stover and Simmonds
1987).
Australimusa is the other section of Musa contributing to edible bananas.
Evolution has led to the entirely different group of edible bananas called Fe’i
banana. It has six recorded wild species, M. lolodensis, M. peekelii, M. maclayi,
M. jackeyii, M. bukensis and M. textilis, while M. fehi is the cultivated species
(Daniells et al. 2001). All are characterized by gigantic stature, red plant sap and
negatively geotropic branching habit (erect bunches). Their distribution is restricted
to New Guinea, Eastern Indonesia, Solomon Islands and other Pacific Islands
(Argent 1976; Nayar 2010). Distribution of Australimusa is well accepted to the
Pacific Islands including Indonesia and the Philippines. But earlier Indian literature
on Sunga dynasty (2 BCE) mentions about a banana variety with red sap. But reports
on banana with red sap raise confusion about the occurrence of Australimusa in
Southern India at some point of time. Archaeological and archaeo-botanical studies
are expected to throw more light on the domestication of Australimusa which started
with the occurrence of parthenocarpy and seed sterility. Mutations and human
selections perpetuated the edible forms especially Fe’i bananas. They were passively
cultivated and protected in many of the west Pacific Islands (Mac Daniels 1947). As
far as evolution is concerned, morphotaxonomic studies of Cheesman (1947), RFLP
markers (Jarret et al. 1992) and Carreel (1994) concluded the interspecific origin of
Fe’i bananas within Australimusa and M. maclayi as the most probable ancestor.
Banana Genetic Resources 323
M. textilis, a member of Australimusa, is endemic to the Philippines. Several

cultivated clones have been evolved in nature as intersectional crosses between M.
textilis and M. schizocarpa and M. textilis and M. balbisiana (e.g. Butuhan, Karoina,
Umbubu, Mayalopa, etc.)
2 Botany and Morphology
Banana is a large, herbaceous member of the family Musaceae belonging to the

order Zingiberales. Earlier it was included under the order Scitamineae.
The true stem of banana is referred as rhizome with roots and vegetative buds.
Simmonds (1962b) called it as corm as it gives a clear-cut idea that banana has erect
underground stem with restricted horizontal growth and possesses roots and
vegetative buds. The centrally located apical meristem gives rise to successions of
leaf primordia which eventually develop into leaf sheath, a strong midrib and an
enlarged lamina. The central stem is a pseudostem formed out of closely packed leaf
sheaths embedding the growing tip.
In general, leaf is uniformly green with various shades except in few cases like
in cvs. Nendran, Pachanadan and other Cavendish clones where purplish
pigmentation is observed on young leaves at early stages. Leaf production rate,
referred as phyllochron, varies from 7 to 10 days under tropical conditions depending
on the prevalent temperature. Appearance and orientation of mature leaves are the
important discriminating traits of various ploidy levels. Diploids have narrow,
slender and erect leaves clustered around the crown. Triploids have normal spreading
leaves, while tetraploids have wider, thick, leathery and drooping leaves. But there
are few exceptions like the Indian cv. Bhimkol, which is a balbisiana diploid having
the appearance and leaf orientation similar to that of a triploid.
The plant puts forth an inflorescence after producing 30–60 leaves depending on
the variety. After the production of all leaves, the plant pushes its inflorescence from
the heart. Inflorescence of banana is a complex spike borne at the end of a stout
peduncle. The flowers are arranged in nodal clusters in two rows, subtended by
spathe like bracts, lanceolate to ovate in shape and pinkish green to deep purple
black in colour. Orientation of the bunch depends on the extent of positive geotropism
expressed by the plant. Based on that, they are referred as follows:
(a) Pendulous – the inflorescence/bunch develops downwards parallel to the pseu-
dostem, e.g. Cavendish clones (AAA) of Eumusa and Ensete glaucum of the
genus Ensete.
(b) Subhorizontal – the bunch is put forth at an angle to the pseudostem, e.g. Silk
and Mysore clones.
324 S. Uma et al.
(c) Horizontal – the bunch is held at almost 90° to the pseudostem, e.g. Ney Poovan
and Pome.
(d) Erect – the bunch looks like a continuation of the pseudostem axis, and the
inflorescence expresses negative geotropism. Here the male bud faces the sky.
Members of Rhodochlamys, Callimusa and, Australimusa sections express this
trait, e.g. M. ornata, M. velutina, etc. of the section Rhodochlamys, M. coccinea
of Callimusa and M. Fei of Australimusa.
Banana flowers are arranged either uni- or biseriately on the nodal clusters on a
nodal cushion. They are covered by spathe like bract. The number of flowers varies
from 1 to 26 depending on the variety. The flowers are of three types, namely, female
flowers, which emerge in the beginning and develop into fruits; hermaphrodite
flowers, which appear immediately after the female phase; and finally the male
flowers. The bracts covering the flowers are an important trait used in identification.
The colour, shape, apex, shoulder ratio, type of lifting, nature of colour fading and
finally the bract scars decide the genomic grouping of an accession through
morphotaxonomic characterization.
3 Taxonomy
Botanically banana is referred to as Musa, and this genus is one among the three
genera in existence, viz. Musa, Ensete and Musella (Liu et al. 2002). They belong
to the order Zingiberales and family Musaceae. The bispecific nomenclature as
Musa sapientum and Musa paradisiaca was suggested by Linnaeus (1753).
Subsequently monospecific nomenclature as M. sapientum ssp. paradisiaca by
Baker (1893), M. paradisiaca ssp. sapientum by Schumann and Engler (1900) and
M. sapidisiaca by Jacob (1952) but none of which were accepted by taxonomists.
Sagot (1887) and Baker (1893) distinguished three subgenera for the genus Musa,
and they were Physocaulis, Eumusa and Rhodochlamys. This classification was
further revised by Cheesman (1947) whose detailed taxonomic classification was
based on chromosome number, pseudostem stature, inflorescence characters and
seed morphology. He divided the genus Musa into four sections, viz. Eumusa
(X = 11), Rhodochlamys (X = 11), Australimusa (X = 10) and Callimusa (X = 10),
and this has been widely accepted. Recently one more section has been added, i.e.
Ingentimusa (X = 7) (Argent 1976; Simmonds and Weatherup 1990).
4 Phylogeny, Domestication and Dispersal
Though there are many species in the section Eumusa, only M. acuminata and M.
balbisiana are believed to have contributed to the evolution of present-day cultivated
bananas which are sterile and parthenocarpic (Stover and Simmonds 1987). India is
well known for its genetic diversity of genus Musa comprising seeded wild species
to seedless cultivars with various levels of ploidy, viz. 2x, 3x, 4x, etc. and different
genomic compositions like AA, AAB, AB, ABB, BB, ABBB, etc. In India, wild
Musa species are largely distributed in the North-Eastern states, the Western and
Eastern Ghats and Andaman and Nicobar Islands.
5 Global Banana Uses, Production and Trade
In India, banana is rightly referred as ‘Kalpatharu’, a plant of all virtues. Besides its
use as dessert fruit and culinary banana, it has multifaceted uses. Banana leaves are
the most popular hygienic dining plate in Southern India. In Africa it is used as
dinner plates and wrapping material, male flower is a much preferred vegetable and
inner core of the pseudostem is also a vegetable in demand with lots of therapeutic
uses. Plant sap is used as indelible ink in industry. Underground rhizome is mostly
exploited as animal feed as a composite mixture with others.
Banana and plantains could be processed into puree, juice, fig, jams and canned
banana slices (Thompson 1995). Banana flour is rich in starch particularly resistant
starch, and this biopolymer constitutes an excellent raw material for baby weaning
foods, puddings, soups, gravies etc. for the preparation of low glycemic products.
Banana wine can be made either from unclarified or clarified juice of beer banana
with a characteristic fruit flavour. A sophisticated distilled alcoholic beverage called
‘waragi’ made from banana is very popular in some African countries like Uganda
and Sudan.
Suitability of banana pseudostem either alone or in combination with paddy
straw produced comparatively higher mushroom yield. They also showed earliness
in spawn and sporocarp production. Banana pseudostem contained substantial
quantity of starch (8.25–8.51%), which could be released mechanically. Banana
juice is a transparent solution which represents 90% of pseudostem weight. The
juice becomes pink when exposed to air and after some time changes to light brown.
The pseudostem when pulped with sodium hydroxide forms black liquor which acts
as an anticorrosive material. The use of spent banana for fibre extraction has gained
momentum across the globe. It has its use in pulp industry, natural water purifier,
bioremediation and recycling, printing Japanese yen, textile industry, as a base
material in cottage industries for making handicrafts and for making a wide range
of goods including rope cordage, yarns, abrasive backing paper, tea bags, shoes, etc.
Commercial alcohol is produced from peel of ripe bananas through saccharification
followed by distillation and fermentation. Yeast of good quality is made from banana
flour, and it can replace malt in breweries. The biowaste obtained at the time of
clearing the land for Jhum cultivation is enormous and easily biodegradable. Hence
it could be vermi composted and used as good manure. Rhodochlamys members
which are short herbs (1–2.5 m height) are highly proliferative in nature with an
326 S. Uma et al.
Table 1 Major banana-producing countries of the world

S. No. Location Countries
1. Latin America and Brazil, Colombia, Costa Rica, Ecuador, Honduras
Caribbean
2. West and Central Cameroon, Côte d’Ivoire, Democratic Republic of the Congo,
Africa Ghana and Nigeria
3. Eastern and Southern Burundi, Kenya, Rwanda, Tanzania, Uganda
Africa
4. Asia and Pacific Australia, Bangladesh, Federal states of Indonesia, India, Papua
New Guinea, the Philippines, Taiwan
average production of 15–20 suckers/year/clump. They have attractive flower buds

coloured with pink, purple, brick red and lilac which could be well exploited in the
cut flower industry.
The major producers of banana namely India, China and Brazil, produce bananas
only to meet out their domestic demand (Table 1). Global banana exports hovering
around 18 million tonnes declined during the past two years due to the spread of
diseases like Fusarium wilt TR4 which negatively affected yield and resulted in
shortages in majority of the producing and exporting regions. Though overall
production increased with an increase in area and productivity, the export demand
is almost static over the years. According to FAO estimates, the major banana
exporting countries are Ecuador, Colombia, Costa Rica and Philippines (Fig. 1) and
major importing countries are the US, Belgium, Germany and UK. The four leading
banana exporting countries in 2015 accounted for 64 per cent of world exports with
Ecuador alone providing more than 30 per cent. The share of the American region
(South, Central America and the Caribbean) decreased from 80 per cent in 1980s to
70 per cent in 2000s. From a continent perspective, Latin American (excluding
Mexico) and Caribbean countries accounted for the highest dollar valueworth of
banana exports with shipments amounting to $7.1 billion or 60.5% of global banana
sales.
5.1 Export Scenario of India
India is the 20th biggest exporter of bananas globally due to the fact that the country
also consumes bananas in greater quantity. India exports US$ 48 million worth
bananas which is roughly 330 crores during the year 2017. Maharashtra almost
monopolized the banana export with the handling volume of 96.91% in value and
the other top banana producing states are catching up significantly for the past three
years. The month wise segregation of data revealed that the peak export was during
April to May (27.6%) whereas in other months, the export is almost stagnant to
6-8%. India export maximum bananas to middle east countries with the lion share
Fig. 1 Banana export by different countries during 2017
Fig. 2 Major Indian banana-importing countries
to United Arab Emirates (41%), followed by Saudi Arabia, Iran, Kuwait and Oman
(Fig. 2). Jawaharlal Nehru Port, Mumbai handles the maximum value of US$16
million (34.7%) followed by Cochin (25.7%), Trivandrum (19.5%), Calicut (8.6%)
through air and through sea port of Tuticorin (6.6%).
328 S. Uma et al.
5.2 B
anana Varieties and Their Distribution
Across the Globe (Table 2)
Banana is called by different names in different languages of the world. The com-
mon names are Pisang in Malaysia and Indonesia, Saging in the Philippines, Kluai
Table 2 Major banana varieties distributed across the globe

Name of
the
S. No. countries Dominant commercial banana cultivars
1. Indonesia Pisang Ambon Kuning (AAA), Pisang Ambon Hijau (AAA), Pisang
Berangan (AAA), Pisang Raja Sereh (AAB), Pisang Mas (AA), Pisang
Berlin (AA), Pisang Kepok (ABB), Pisang Raja Bulu (AAB), Pisang Tanduk
(AAB), Pisang uli, Pisang Mas, Pisang senm, Pisang Kosta, Pisang Kosta
putih, Pisang kepo, Pisang Lampung, Pisang Jari Buaya, Pisang Nangka
2. Malaysia Pisang Berangan (AAA), Pisang Mas (AA), Pisang Cavendish (AAA),
Pisang Rastali (AAB), Pisang Tanduk, Pisang Raja (AAB), Pisang Nipah
(AAB), Pisang Awak (ABB), Pisang Lemak Manis (AAB), Pisang Nangka
(AAB), Pisang Mas, Pisang Embun, Pisang Mask Hijau, Pisang Rejang,
Pisang Keling
3. Brazil Prata (Pome) (AAB), Pacovan (AAA), Prata Ana (AAB), Maca (Silk)
(AAB), Mysore (AAB), Nanica (Cavendish) (AAA), Naniciio (Cavendish)
(AAA), Terra (Plantain) (AAB), Anglao (Plantain) (AAB)
4. Philippines Buñgulan (ABB), Cardaba (ABB), Saba (ABB), Grand Naine (AAA),
Cuarenta Dias (AAA), Lagkitan (AAA), Lakatan (AAA), Latundan (AAA),
Cachaco (ABB), Gros Michel (AAA), Pisang Ceylan (AAB), Pisang Jari
Buaya (AA), Sukali Ndizi (AA), Bogoya (AAB), Gonja (AAB), Kayinja
(ABB), Kisubi (ABB), East African Highland banana (EAHB), Inaribal
Katisla, Pelipita
5. Thailand Klui Khai (ABB), Klui Le Mu Nang (AA), Klui Thang ruang (AAB), Klui
Khom Thonng, Klui Hom Kneio, Klui Hom Khom, Klui Nang, (ABB), Klui
Lacka, Klui Namwa Khom (ABB), Klui Lep Chang Kut (BBB), Klui Khi
Bong (ABB), Klui Hak Muk Kho, Klui Muok Khieo, Klui Teparot (ABB)
6. Vietnam Chuoi Tay But, Chuoi Nguothoc, Choui Tieululn, Chuoi Tieu Vue, Chuoi
Tieu Cao, Thoui, Man, Chuoi Tay, Chuoi Ngoptun, Chuoiomat, Choui Mat
boket, Chouii Tieincuoioui
7. India Robusta (AAA), Red Banana (AAA), Matti (AA), Anaikomban (AA),
Hanuman (AAA), Thella Chakkarakeli (AAA), Amrit Sagar (AAA), Dwarf
Cavendish (AAA), Kunnan (AB), Ney Poovan (AB), Poovillachundan (AB),
Chevvazhai (AAA), Manoranjitham (AAA), Monthan (ABB), Karpuravalli
(ABB), Nendran (AAB), Nedunendran (AAB), Chengalikodan (AAB),
Rasthali (AAB), Mortman (AAB), Poovan (AAB), Mysore Bale (AAB),
Kadali (AA), Hill banana (AAB), Sirumalai (AAB), Virupakshi (AAB), Ladan
(AAB), Nadan (AAB), Jwaribale (AAB), Nendrapadathi (AAB), Chinia
(ABB), Peyan (ABB), Pisang Rajah (AAB), Thenkadali (ABB), Boddida
Bukkisa (ABB), Bangrier (ABB), Vayalvazhai (ABB), Chakkiya (ABB)
8. Uganda East African Highland banana (EAHB), Sukali Ndizi (AA), Gonja (AAB),
Kayinja (ABB), Kisubi (ABB),
9. Costa Rica Gran Enano, Valery
10. Colombia Gros Michel and Pátano Enano
in Thailand, Choui in Vietnam, Kela/Vazhai/Vazha/Bale and Arati in India and

Chiao in China which are applicable to all dessert and cooking bananas, including
plantains.
6 Status of Wild Musa in India and Their Distribution
Musa species was first reported in India by Hooker as early as 1892. In India, more
than 11 species have been reported including M. acuminata ssp. burmannica,
M. acuminata ssp. burmaniccoides, M. sikkimensis, M. balbisiana, M. nagensium,
M. thomsonii, M. itinerans, M. ochracea, M. flaviflora, etc., which are widely
distributed in the Northeastern India (Uma et al., 2001). M. balbisiana is the other
wild Eumusa species which had fairly broad distribution across South and SE Asia.
It is fairly a stable species without much variability within unlike M. acuminata. But
recent explorations and molecular characterization have revealed sufficient
variability in stature of the plant, pseudostem colour, leaf orientation, male flower
bud shape, etc. which may result in few subspecies (Uma et al. 2005; Ge et al. 2005;
Sotto and Rabara 2000). Besides Eumusa, Rhodochlamys are also abundantly
distributed in the states of Northeastern India and Western and Eastern Ghats.
M. ornata is distributed in Western Ghats and Eastern Ghats, whereas the species
like M. velutina, M. aurantiaca and M. rosacea are commonly growing in the undis-
turbed forest areas of Northeastern India. M. laterita, a unique species which is very
short with rhizomatous roots, is found in Western Ghats (Abraham 1976). Regarding
Callimusa and Australimusa, no records are available for the presence of members
belonging to these two sections in India.
Recently some more new species have been reported in India: M. swarnaphalya
(Uma et al. 2005), M. saddlensis and M. kuppiana (Anon. 2005), M. velutina subsp.
markkuana (Sabu et al. 2013), M. velutina var. variegata (Joe et al. 2013b) and M.
sabuana (Prasad et al. 2013), M. nagalandiana (Dey et al. 2014), Musa balbisiana
var. andamanica (Singh et al. 1998) and M. arunachalensis (Sreejith et al. 2013)
(Tables 3, 4, and 5).
7 Diploid Bananas
Wild bananas are all diploids, and the ancestry of cultivated bananas revealed the
involvement of two major species M. acuminata and M. balbisiana with occasional
mention of M. textilis and M. schizocarpa. Simmonds (1962b) proposed early
domestication of diploid banana, i.e. M. acuminata in its primary centre of origin,
namely, Malaysia, with its large variability even today. The evolution from wild to
cultivated bananas involved seed suppression and development of parthenocarpy
(Perrier et al. 2011). The partial occurrence of parthenocarpy in M. acuminata ssp.
banksii (Simmonds 1962a, b) might have contributed for the development of
330 S. Uma et al.
Table 3 Distribution of wild Musa species in India

S. No. Name of the species Distribution
1 M. acuminata ssp. Western Ghats
burmannica
2 M. acuminata ssp. Western Ghats and Northeastern India
burmaniccoides
3 M. balbisiana Western Ghats, Andaman Nicobar Islands and
Northeastern India
4 M. itinerans Arunachal Pradesh, Nagaland
5 M. cheesmanii Arunachal Pradesh, Meghalaya and Nagaland
6 M. ochracea Tripura, Manipur and Arunachal Pradesh
7 M. flaviflora Arunachal Pradesh, Assam, Tripura, Mizoram,
Meghalaya and Manipur
8 M. sikkimensis Arunachal Pradesh, Nagaland, Manipur, Tripura and
Meghalaya
9 M. nagensium Nagaland and Mizoram
10 M. thomsonii Nagaland and Mizoram
Table 4 Distribution of recently identified Musa species in India

1 M. swarnaphalya Arunachal Pradesh (Uma et al. 2011)
2 M. saddlensis Arunachal Pradesh (Anon. 2005)
3 M. kuppiana Arunachal Pradesh (Anon. 2005)
4 M. velutina subsp. markkuana Arunachal Pradesh (Sabu et al. 2013)
5 M. velutina var. variegata Arunachal Pradesh (Joe et al. 2013b)
6 M. sabuana Arunachal Pradesh (Prasad et al. 2013)
7 M. nagalandiana Nagaland and Manipur (Dey et al. 2014)
8 M. balbisiana var. andamanica Andaman and Nicobar Island (Singh et al.
1998)
9 Musa arunachalensis Arunachal Pradesh (Sreejith et al. 2013)
Table 5 Distribution of Rhodochlamys and allied genus Ensete in India

1 M. laterita Western Ghats and Assam
2 M. aurantiaca Arunachal Pradesh
3 M. velutina Assam, Arunachal Pradesh and Meghalaya
4 M. ornata Tamil Nadu, Andhra Pradesh, Mizoram
5 M. rosaceae Arunachal Pradesh
6 Ensete superbum Western Ghats of Kerala, Karnataka, Maharashtra and Gujarat
7 Ensete glaucum Northeastern India
parthenocarpy which was later confirmed by RFLP studies (Carreel 1994).

Simultaneous development of parthenocarpy has been reported by Uma et al. (2005)
while studying M. acuminata ssp. burmannica having distribution in Western Ghats
of India and Matti, parthenocarpic landrace using morpho-molecular characterization.
Matti is suggested to have evolved due to continuous human selection for complete
parthenocarpy likewise Pisang Rejang and the possible seedy progenitor of Pisang
Lilin (Nasution 1991); a parthenocarpic diploid in Indonesia also confirms the
simultaneous and independent development of parthenocarpy in banana.
Unlike M. acuminata, the development of parthenocarpy in M. balbisiana has
not been proved scientifically. Occurrence of parthenocarpic diploid and triploid M.
balbisiana (BB and BBB) has been reported from the Philippines and Thailand by
several workers, but their true genomic constitution still needs confirmation.
Apomictic seed development in Ensete superbum induced by pollinating with M.
balbisiana (W) pollen has been reported by Ravishankar et al. (2011). Some
accessions reported to be BBB in the gene bank at Phakchong Banana Research
station, Thailand, were found to exhibit the traits of M. textilis. A comprehensive
study using morphological and molecular approaches is warranted to decipher the
actual genomic status. But edibility in terms of emptiness of seeds, softened seed
coats and pulpiness of fruits over its mucilaginous pulp has been observed in
‘Bhimkol’, a widespread landrace of Northern Eastern India.
It appears that parthenocarpy in banana is genetically controlled by one major
gene and two minor modifier genes (Ortiz 1995; Ortiz and Vuylsteke 1995 and
Okoro et al. 2011). Sterility is also genetically manoeuvred, supplemented by
various forms of chromosomal aberrations and abnormal meiosis, all contributing to
edibility in banana. Vegetative propagation favoured the perpetuation of edibility.
Human intervention for selection of edible diploids and nature’s intervention to
induce sterile mutants resulted in vast diversity of AA diploids which are manifested
as cultivated diploids. Another major milestone was the development of better
edibility in terms of sweetness. M. acuminata ssp. banksii-derived diploids had
starchy fruits, and sweet cultivars are suggested close to M. acuminata ssp.
malaccensis (Carreel et al. 1993). Similarly clones from the different subspecies
differed by translocation events on their chromosomes (Shepherd 1999). Natural
introgression between clones of subspecies and human selection for better edible
diploids was the vital step in banana evolution. Presently large variability is seen in
South and Southeast Asia including the Pacific Islands.
8 Polyploid Bananas (Triploids and Tetraploids)
By virtue of their superior traits like parthenocarpy, edible pulp, good bunch and
sturdiness, triploids were preferred for selection and clonal perpetuation by early
humans. Female restitution leading to the production of diploid gametes and
hybridization among the subspecies of M. acuminata or different species led to the
production of triploids. It is reported that production of unreduced diploid gametes
332 S. Uma et al.
is quite frequent in both diploid males and females (Dodds and Simmonds 1946).
Naturally occurring improved edible diploids of a particular subspecies, retaining
fertility to some extent and hybridizing with other subspecies, might have led to the
development of initial acuminata triploids. But tetraploid acuminata (AAAA) is not
a commonly reported phenomenon in nature.
8.1 Bispecific Polyploids
Introgression of acuminata diploids with M. balbisiana occurred either in their pri-

mary origin or little away on their journey to new locations resulting in interspecific
crosses. This resulted in different genomic combinations, viz. AB, AAB, ABB and
ABBB of which AAB and ABB encompassed a greater diversity for modern
bananas.
AAA genomic group forms the major allopolyploids valued for its commercial
status in global banana industry. It has various subgroups like Cavendish, Gros
Michel, Red, Ibota, Mutika/Lijugera and a couple of unique accessions (Uma and
Sathiamoorthy 2002). Cavendish clones represent more than 50% of global banana
industry and are expanding in terms of area and value by virtue of their high
productivity and resistance to Fusarium wilt race 1 and 2. Gros Michel was the
earlier ruling variety replaced by Cavendish. Red group has not much variability
except for its green mutant. Mutika/Lijugera comprises East African Highland
bananas contributing to staple food in Rift Valley region of Burundi, Rwanda and
Uganda. They are cooked and brewed for beer.
AB diploids evolved in specific locality of SE Asia especially Southern India and
Sri Lanka (Uma and Sathiamoorthy 2002) from where it got distributed to Eastern
Africa in the recent past (Shepherd 1957; De Langhe 1996; De Langhe and De
Maret 1999; Nayar 2010). Indian classification of AB genome has Ney Poovan and
Kunnan as its subgroups, while the international classification system (Daniells
et al. 2001) has Ney Poovan and Kamarangasenge. Molecular-assisted diversity
analysis (SSR) confirmed the occurrence of only two subgroups in the AB genome,
Kunnan and Ney Poovan. Ney Poovan has minimum morpho-molecular diversity,
while Kunnan exhibited higher phenotypic diversity (Uma et al. 2010). The study of
AB genotypes specific to South India, viz. Kunnan and Safed Velchi, for their
lineage, gave no evidence of balbisiana cytoplasmic genome (Carreel et al. 2002).
It is possible that the M. balbisiana types used in the study are SE Asia origin which
could be different from those of Indian origin. India has diversity for M. balbisiana
(Uma et al. 2005, 2006a, b). Inclusion of diversity from India for such studies is
expected to provide a holistic picture on its evolution.
8.2 AAB Cultivars
Introgression between two species in various combinations, especially M. acumi-

nata ssp., led to a broader diversity classified as Plantain, Silk, Mysore, Pome and
other unique members. Evolution of plantains from M. acuminata ssp. banksii as
female parent and M. balbisiana as male parent has been well demonstrated by
maternal and paternal lineage studies using RFLP (Carreel et al. 2002). This sug-
gests that plantains evolved in New Guinea where a greater diversity for plantains
and cultivated diploids exist naturally. Southern India could only be a secondary
centre of diversification as it has only French plantain.
Silk group of bananas are popular cultivars of South and Southeast Asia, East
Africa, Brazil and other Latin American countries. Maternal paternal lineage studies
using RFLP (Carreel et al. 2002) reveal its paternal lineage from M. acuminata ssp.
malaccensis unlike other AAB subgroups. Subspecies malaccensis has natural
distribution in Malaysia, Indonesia and Thailand where primarily evolved diploids
introgressed with M. balbisiana in nature to give rise to Silk clones (AAB).
Mysore group (AAB) is another group of hardy bananas spread across the world.
Evolutionary studies revealed M. acuminata ssp. errans as its lineage and M.
acuminata subspecies as maternal with cp DNA II (Carreel et al. 2002). This also
has AB diploid and M. balbisiana in its ancestry. Subspecies errans has its
distribution mainly in the Philippines. But the natural occurrence of M. swarnaphalya,
which is morphologically akin to M. acuminata ssp. banksii or errans in Northeastern
India, could lead to another centre of evolution for M. acuminata ssp. banksii and
M. acuminata ssp. errans-derived bananas. Pisang Kelat, a unique AAB member, is
reported to have the maternal origin of the B genome and hence classified as
BAA. This lineage is important from pathological perspective since this is an
accession immune to most of the leaf spot diseases and Fusarium wilt (Anon. 1999).
Next important group is Maia Maoli/Popoulu (AAB), which are endemic to
Oceania and specifically of Polynesian origin. Maoli/Popoulu group is recognized
for unusually fat and squat fruits with blunt tips, but Maoli group exhibits longer
fruits. Lot of diversity for fruit shape and size exists in Hawaii, West Polynesia,
Pohnpei, etc. Popoulou fruits are very characteristic due to their tendency to split at
ripeness (Ploetz et al. 2007).
Iholena subgroup is traditionally grown in highlands of Hawaii distinguished by
coppery under shades of leaves, Salmon-coloured flesh and right-angled fruits.
They are consumed raw or cooked. Hawaiian diversity is represented by seven
Iholena varieties. Iholena fruits are arranged loosely and at right angles to the bunch.
Male flowers have characteristic lavender-coloured stamens. Fruits remain pale
yellow green throughout its development. Iholena is also sparsely represented in
New Guinea, Samoa, French Polynesia, Tonga and others (Ploetz et al. 2007). The
uniqueness of the group and its diversity are attributed to a series of additional
somatic mutations in basic hybrids and cultivars in niche locations of Polynesia (De
Langhe 1996). Their genetic closeness to plantains is well evidenced by Carreel
334 S. Uma et al.
et al. (1993) with M. acuminata ssp. banksii contributing to AA genomes in both

cases (Horry and Jay 1988; Lebot et al. 1993; Carreel et al. 2002).
8.3 ABB Cultivars
All ABB clones in various subgroups of Bluggoe, Pelipita and Ney Mannan have
lineage of Musa balbisiana (Carreel et al. 2002) and may be referred as true ABB,
while Pisang Awak and Peyan, the dessert cultivars of ABB, have cytoplasmic DNA
associated with B genome and are referred as BAB. International classification of
Pisang Awak and Peyan (dessert bananas) and Monthan, Bluggoe and Vennutu
Mannan (cooking types) has been suggested to include Bontha as another subgroup
which is being debated. Most of them are starchy, primarily cooked and eaten. Some
Bluggoe types are commercially used as ripe fruits in South India. Though it is said
that B-rich genomes are drought tolerant, Monthan and others are few exceptions.
9 Status of Cultivated Bananas in India
India is one of the countries with different agroclimatic conditions which have
encouraged the development of different and large number of varieties catering to
local needs. Even though India is having vast diversity for banana and plantains,
only few are cultivated commercially in many states of India, and banana trade is
dominated by only one or two cultivars especially Cavendish type (Grand Naine). It
is known that 15–20 cultivars are grown based upon the local needs and consumer
preferences. Common banana cultivars and states grown are given in Table 6. Apart
from the common commercial cultivars, some other cultivars are also available, but
their utilization is meagre. Those less exploited banana landraces were identified
and documented (Uma et al. 2014).
10 Evolution of Ensete
Ensete is the least studied genera in the Musaceae with little work on their origin
and evolution. But it is suggested that both Ensete and Musa diverged from a
common stock (Cheesman 1947; Simmonds 1962b; Chakravorti 1955), but, not
sensu stricto, is derivable from the other. Occurrence of M. ventricosum in North
America (middle Eocene of Oregon) about 43 million years ago has been reported
by Manchester and Kress (1993). Evidence of domestication has been well reported
since prehistoric period (Brandt 1984; Clark 1967). During the course of evolution,
Ensete are reported to have retained most of the primitive characters. Chakravorti
(1955) proposed the evolution of Eumusa with 2n = 2x = 22 from Ensete,
Table 6 Major banana-growing states and the common cultivars

S. No. State Common cultivars
1 Tamil Nadu Matti (AA), Grand Naine (AAA),
Robusta (AAA), Red Banana (AAA),
Dwarf Cavendish (AAA), Manoranjitham (AAA), Ney Poovan (AB),
Poovillachundan (AB), Nendran (AAB), Poovan (AAB), Pachanadan
(AAB), Rasthali (AAB), Hill banana (AAB), Chakkiya (ABB),
Vayalvazhai (ABB), Karupuravalli (ABB),
Monthan (ABB) and Peyan (ABB)
2 Kerala Grand Naine (AAA), Robusta (AAA),
Dwarf Cavendish (AAA), Red Banana(AAA),
Ney Poovan (AB), Kunnan (AB), Nendran (AAB), Poovan (AAB),
Rasthali (AAB),
Chakkiya (ABB), Karupuravalli (ABB) and
Monthan (ABB)
3 Andhra Grand Naine (AAA), Robusta (AAA),
Pradesh Red Banana (AAA), Thellachakkarakeli (AAA), Ney Poovan(AB),
Karpura Chakkarakeli (AAB),
Amrithapani (AAB), Boddida Bukkisa (ABB),
Monthan (ABB)
4 Bihar Grand Naine (AAA), Bhusawal (AAA),
Lalkela (AAA), Ney Poovan (AB),
Champa (AAB), Malbhog (AAB),
Kanthali (ABB), Monthan (ABB)
5 West Bengal Grand Naine (AAA), Bhusawal (AAA),
Lalkela (AAA), Ney Poovan (AB),
Champa (AAB), Malbhog (AAB),
Kanthali (ABB), Monthan (ABB)
6 Karnataka Grand Naine (AAA), Robusta (AAA),
Chandrabale (AAA), Dwarf Cavendish (AAA),
Elakkie Bale (AB), Mysore Bale (AAB),
Rasabale (AAB), Bargi Bale (AAB), Nanjangud Rasabale (AAB),
Sarubale (ABB), Sakkarbale (ABB)
7 Maharashtra Grand Naine (AAA), Safed Velchi, Mysore Ethan
8 Gujarat Grand Naine (AAA)
9 Assam Jahaji (AAA), Amrit Sagar (AAA), Malbhog (AAB), Cheni Champa
(AAB), Kachkel (ABB)
10 Tripura Jahaji (AAA), Amrit Sagar (AAA), Malbhog (AAB), Cheni Champa
(AAB), Kachkel (ABB), Rigatchi (ABB)
11 Odisha Patkapura (AAB), Bhusawal (AAA), Buntal (ABB)
2n = 2x = 10, which earlier referred as genus Physocaulis. Twenty-two chromo-
somes of Eumusa were derived through fragmentation of two particular chromo-
somes across their secondary constrictions followed by recombination of the
resulting fragments. He supported his theory based on close karyotypic resem-
blances between M. acuminata and M. superbum (presently E. superbum). Although
not much information is available, recent explorations in NE India have led to the
identification of a new species, tentatively named as M. kuppiana, with intermediary
336 S. Uma et al.
morphological traits of Musa and Ensete (Anon. 2006). Like plantains, origin of
Ensete is accepted to be South and Southeast Asia and then spread to Africa where
it got diversified into several species with great adaptations suitable for cultivation
at altitudes ranging from 600 to 3000 m.
11 Exploration and Collection of Musa Genetic Resources
Musa germplasm collection is one of the primary activities of plant genetic resources
centres. It aims at capturing plant genetic diversity and, in this context, is an initial
step to back up all the related PGR activities. The growth and development of this
discipline and the activities taken up in India are highlighted in this section. The
theoretical know-how and its implications against ethnic and cultural background of
farmers who grow landraces are discussed and suggestions given to make collecting
of germplasm more practicable. It is advocated that the discipline, being field-based,
involves considerable knowledge of plant geography, agroecology, plant taxonomy
studies, ethnobotany, crop evolution and domestication, population variation and
distribution, gene pool sampling and several other related topics. It is clear that the
only safe approach to provide broad genetic bases and satisfy the needs of future
plant breeding programmes is to collect and maintain as much as possible of the
entire genetic diversity of both cultivated species and their wild relatives.
Collection of banana genetic resources primarily aims to assemble as much vari-
ability as possible, that is, available in the cultivated banana and its wild relatives.
The germplasm so collected reveals the nature and extent of variability in different
genotypes, cultivars, etc. and their agroecological/phytogeographical distribution.
The field sampling procedures in Musa genetic resources exploration are aimed at
the fullest possible recovery of genetic variation available within genus/species/
cultivars/landraces. Strategic planning is very much required prior to taking up of
exploration and collection so that the explorer is in the right area at the right time to
collect his target germplasm such as suckers/vegetative propagules, seeds, etc. and
study the existing variability in the natural habitat. Knowledge on agroecology,
crops, stage and their distribution in the areas of survey, local contacts, equipment
required, transport arrangements and routes to be followed, distances involved,
places of halt/camping sites available, transport of material, team composition, etc.
is to be acquired before setting out on a collecting expedition.
Musa germplasm collecting missions are the efforts of understanding the preva-
lent genetic diversity in different areas. A germplasm collection can include wild
relatives of domesticated species, wild species directly used by human beings, obso-
lete cultivars, advanced cultivars, landraces and breeding lines.
It is necessary to acquire knowledge on the spectrum of genetic diversity occur-
ring in the centre of diversity and in the areas of its cultivation. Germplasm collec-
tion involves application of both the theoretical knowledge on population sampling
and practical know-how in overall understanding of plant diversity and environment
including the socio-economic and cultural aspects of the farming societies.
12 Technical Guidelines for Exploration
Exploration planning involves prior knowledge of the area where the exploration is
to be made, its people (cultural communities, ethnic groups, tribal, etc.), socio-
religious customs, eco-edaphic conditions, genus/species/landraces grown and the
varietal diversity available. Before sketching out an expedition programme, an
explorer must gather all the available information which would make it into a
successful venture (Frankel and Soulé 1981; Hawkes 1983; Arora 1991):
1 . Deciding the route of expedition and sites for collection
2. Strategy of gene pool sampling
3. Equipment to be carried for collection and transport of material
4. Other miscellaneous prerequisites
Further, soundness of the political climate of the terrain to be explored is also of
primary concern while planning germplasm collecting missions. These aspects are
briefly dealt below.
Acquiring Knowledge on Agroecology and Crop Distribution
It would be necessary to acquire knowledge on the agroclimatic conditions in
relation to the distribution of species/varieties/taxa of crop(s) and/or their wild
relatives in the area to be explored. In this context, state reports, regional documents,
floras, floristic surveys and other published works would be of great help to
familiarize the plant explorer with the climate, ecology, vegetation and agriculture.
More emphasis should be laid on latest published/unpublished reports of the plant
explorers/surveys carried out. Such literature would provide clues to flowering and
fruiting states of Musa and its allied species available in relation to agroecology of
the terrain. Local habitat variations within an agroecological region are often
enormous and so also the ecotypic variations.
Visit to Genetic Resources Centres
It would be desirable for an explorer to plan a visit to the genetic resources centre
or institute/other crop-based institute of relevance to his mission. If planned in the
crop season, he can see (part of) the collections in the field. At least, the resident/
country collectors must get familiarized with such collections. When the collector is
directed to collect specific materials possessing specific attributes, discussions with
crop breeders/botanists maintaining and evaluating local/regional/national
collections provide useful clues to the areas of survey. Published articles and
catalogues on crops also provide such useful information.
Establishing Local Contacts
It is necessary to list out the local agencies and/or centres in the area of survey
and establish administrative/scientific contacts, with State Agricultural Departments,
Block Development Offices and their branches and Regional/National Institutes.
338 S. Uma et al.
Information on the locations proposed to be visited, routes to be followed, position

regarding camping sites, distances between places en route, location of petrol
pumps, crops and their harvesting time should be discussed with local counterpart(s)
before embarking on the mission. Also, it would be better to develop contacts with
scientists engaged in crop-specific research programmes and to correspond with
local contacts in order to know more about the nature of variability available in
native genetic resources – landraces and their distribution, occurrence of wild Musa
species, etc.
Planning of Itinerary
Information collected as above would help in the planning of proper itinerary. A
provisional route and time schedule can be worked out based on the harvesting time,
agroecology of the terrain and the distribution of crop diversity, wild relatives,
endemic types and accessibility to the area, etc.
12.1 Logistic Preparation/Implementation
Larger teams usually prove troublesome, while small teams are more effective and
mobile. A two-member team is considered ideal with a local helper and crop
specialist/extension worker. Smaller teams are welcomed and can build up
confidence quickly through the local officer/guide. The farmers of underdeveloped
tracts holding native landraces do not like or permit the team to visit their fields, and
the smaller the party, the lesser the interference and the more the possibility to
survey such sites. In some specific exploration missions, larger teams are
unavoidable. Here, the mission would have a crop scientist/plant pathologist, and it
is an additional advantage if one such member knows driving also.
Team Composition
When the team has more members, the leader must coordinate collecting activi-
ties right from the initial planning. Sometimes, team members are from different
centres/organizations, and an effective contact is to be maintained with them by the
team leader. Before the collecting programme starts, the leader must thoroughly
brief the party members in the first meeting about the objective of their mission.
Duration of Exploration
This would vary according to the mission. Explorations within the country are of
shorter duration; 4 weeks or even less would be ideal and recommended, as
otherwise exploration missions tend to be more expensive and less purposeful.
Basic Field Exploration Equipments (Table 7)
Above all, jeep is ideal for a small party, and a covered van is also good, prefer-
ably with improvised sleeping arrangement. Four-wheel-driven car with a collaps-
ible roof tent on top of the vehicle is ideal for small teams (two or even three
Table 7 Important items and equipments required

Survey/ (a) Altimeter (with adequate altitudinal range), compass, soil kit, SLR camera
collecting (35 mm), binoculars, hand lens, GPS, different colour cloths for taking
items photographs of various plant parts
(b) Haversack/kitbag, cloth bags, alkathene bags, aluminium labels, drying
sheets, old newspaper, plant press, vasculum, rubber bands, gum tape, rope
(thick and thin), scissors, knife, trowel and digger and for field data recording:
field notebook, collecting sheets, diary, pencil, ball pen, red/blue pencil/marker,
stapler, etc.
Published Regional flora, other reports, list of local names of crop plants/cultivars; road
material map, vegetation/climate map, list of rest houses/lodges, hotels, resting/stay
places, petrol points (diesel, petrol), distances between sites/towns en route,
jeepable tracts, fair weather roads, etc.
Medicines, Antimalarial pills, APC tablets, anti-amoebic and antidiarrhoeal tablets, Vicks,
First Aid kit, Odomos, Burnol, antiseptic cream/Savlon/Dettol, cotton packs, Johnson
etc. band-aid, furacin powder, dressing gauze, water-purifying tablets, first-aid box,
etc.
Other Water bottle, torch (with extra battery cells), large candles, match boxes, steel
equipments box, cardboard/carton boxes or other containers, printed slips with the
institute’s address, plastic jars, formaldehyde, alcohol, hunter shoes, tarpaulin,
camp cots, sleeping bags and cooking items depending on need, cap for sun
protection, etc.
persons). Take at least two extra jerricanes, one or preferably two spare tyres and
spare parts like fan belt.
12.2 Practical Guidelines
Having acquired the basic knowledge on the aspects of sampling, practical consider-
ations would demand application of certain other tactics, as discussed below.
12.2.1 Survey Tactics
1. Depending on the area of survey, the explorer should cover drier sites earlier and
the humid belts later. Likewise, unirrigated pockets holding primitive germplasm
would need survey much ahead of that to be conducted in the irrigated terrain.
Also, in the hills, normally lower valleys need to be covered earlier than the
higher elevated areas, barring special ecological situations such as cold arid des-
ert in the Western Himalayas.
2. The mandate of the exploration team is to locate types specifically adapted to
particular environments/sites, which vary in soil and climate and topography/
elevation, and visualize climatic variation vis-a-vis itinerary properly so as to
collect specific types suited to such edaphic ecological situations and
physiologically stress situations (saline habitat, grown under unirrigated
condition/drought adaptable and cold adaptable). It is advantageous to collect
340 S. Uma et al.
germplasm in sites/villages much away from the approachable roads, as the

easily accessible sites must have been previously explored. Collection must be
planned in inaccessible pocket valley and isolated hills, difficult to approach
even on foot. Situations representing limits of agriculture, namely, altitudinal
limits of agriculture in mountains or deserts, may offer different germplasm.
Likewise, villages at the edge of the desert, isolated coastal belts (islands) and
the ecotonal belts in case of wild germplasm would provide more exactly the
sites of potential diversity.
The areas holding landraces are invariably dominated by native people who eke
out marginal existence based on multi-crop subsistence agriculture practised on
small landholdings. Often it is observed that the natives consume the previous year’s
harvest. So, only limited/meagre collection will be possible when exploration is
organized in such areas during seasons other than the crop maturity period. In such
situations, the plant explorer should stress on collecting from the previous year’s
harvest and take care not to collect infested seeds.
The collector must keep in mind that crops often vary with ethnic diversity, and
different arrays of materials may be collected even from contiguous belts occupied
by different tribes. The explorer should survey such sites/villages very thoroughly,
as invariably such locations are rich in materials in which endemic characters are
numerous. Here, the village (tribal) chief is a very important contact person. Local
guide, or interpreter accompanying the party, can bridge the gap and create goodwill
and confidence. Further, it is advocated that in such remote survey areas, the team
may carry token gifts for the natives. Occasionally, some token money may have to
be paid to these farmers to get their harvested produce so as to collect the desired
germplasm. But this must be negotiated first through local counterpart/guide and
implemented only when permitted by the village chief, who would be then the first
one to receive such gift items.
Transportation of Material
If the team is to send plant/germplasm collections to the institute, halting in
between at a bigger town may be necessary. Incidentally, this halt can also be
utilized for re-equipping provisions, jeep servicing/checkup and purchase of several
other articles needed by the team. For long-duration explorations in other countries,
transportation of collections to destination would be required, and halts at towns
with air-transport post-dispatch facility, etc. are advocated, though as per
prearrangement, normally collections would need phytosanitary clearance from the
authorized agency in that country.
Funding Aspect
Exploration funding is always worked out tentatively. The team occasionally
may find this calculation as an underestimate. Additional funds may be required. So,
keeping track of accounts is essential, and the team leader is to be vigilant about
fund position. If more funding is needed, take action much ahead of time using
facsimile, telex, telephone or cable – a quick message delivering facility through the
local government. One has to pre-plan this action so that the work is not held up. In
general, include unforeseen expenses while estimating funding of expeditions. Keep
enough local currency for use during exploration – en route travel in small towns,
money of smaller denominations and coins for village transactions.
After Expedition Care
Follow-up would be needed so that the accessions collected are kept safe, index-
ing/accessioning may be done and collections passed on for quarantine. Be sure that
the field books and survey/field notes are complete for writing the final report. The
use of a portable computer containing the collecting sheet format is becoming more
and more a common practice and can be a considerable time-saver.
12.3 Knowledge on Agroecology and Musa Distribution
It is necessary to acquire knowledge on the agroclimatic conditions in relation to the

distribution of species/varieties/wild relatives in the area to be explored. In this
context, reports of state forest departments/horticultural and agricultural department/
Botanical Survey of India (BSI) and other published works would help a lot to
familiarize the plant explorer with the climate, ecology, vegetation and agriculture.
More emphasis should be laid on latest published/unpublished reports of the plant
explorers/surveys carried out. Such literature would provide clues to the harvesting
time of the crops, prevalent agri-horticultural diversity and the economic plants
available in relation to agroecology of the terrain. Local habitat variations within an
agroecological region are often enormous and so also ecotype variations. GIS and
ecogeography have several applications in collection, conservation and efficient use
of banana genetic resources. In recent past, their use in conservation and sustainable
use of agro-biodiversity is increasing considerably to counteract the challenges
faced in the context of global climate change (Jarvis et al. 2008). These advanced
tools and studies are very much essential in banana to identify the most suitable site
for the regeneration of a wild germplasm which has potential use in breeding of
improved varieties with resistance to biotic and abiotic stresses. They also enable us
to strategically plan for in situ conservation which in turn might reduce the genetic
erosion of a species or loss of specific traits due to its non-adaptation in a specific
environment.
12.4 In Situ Observations
Data at the collection sites are very important, and they must be documented neces-
sarily for future references. The details on the data to be taken are as follows:
1. Passport information/data
342 S. Uma et al.
2 . Minimum description of the species/landrace/variety

3. Documentation in the form of photographs
4. Traditional/indigenous knowledge, wherever applicable
The amount of information which can be recorded during collecting activities is
very much dependent upon the location, situation at the site and time available.
Information of primary importance must be recorded at the place of collection
itself. But other details regarded as of secondary importance should be kept at
minimum so that the time spent on collecting can be maximized. Different types of
collecting forms, sheets or books have been used by collecting missions. Usually
Musa explorers are being used for recording collection site information and plant
description, passport data book (NBPGR), Musa descriptors (IPGRI/INIBAP
1996) and minimum descriptors developed by NRCB (Anon. 2015). Such docu-
mentation of data on genetic resources has been compiled in the form of an ‘Indian
Banana – Genetic Resources Catalogue’ by ICAR-NRCB, Trichy (Uma et al.
2005). The data is updated periodically in MGIS providing access to the banana
scientific community.
The Musa Germplasm Information System (MGIS) is a user-friendly database
which contains key information on Musa germplasm diversity, including passport
data, botanical classification, morphotaxonomic descriptors, molecular studies,
plant photographs and GIS information on 4589 accessions managed in 21
collections around the world, making it the most extensive source of information on
banana genetic resources. This database could also be used for the identification of
an unknown banana genetic resource. The database is managed by Bioversity
International office at Montpellier, France.
12.5 Germplasm Collection
Sampling Strategy
The sampling strategy largely depends on the crop species (self- or cross-pollinated
or vegetatively propagated) and the extent of gene exchange between populations,
so also on the primary objectives of collecting the genetic diversity in the area being
covered.
Though the theory of sampling strategy stresses upon extensive knowledge of the
patterns of genetic variability of populations, in general, there are relatively few
species for which this type of information is available. Most species exhibit extensive
geographical variation, and superimposed is the variation within populations.
Ecological habitat and/or factors are a major determinant of genetic diversity, and
agro-ecotypes are most clearly distinguished from primitive cultivars and landraces.
It is also realized that geographical variation patterns include such characteristics as
disease resistance, morphological features and other conspicuous differences as
well as variation in quantitative characters relevant to plant breeders. Though
varieties or strains may look alike, they would differ greatly in useful attributes,
especially in physiological characters. Sampling methods used must ensure the

collection of representatives for the variation within population as well as those
associated with geographical patterns of variation.
The principles in selecting the site (field) and sample numbers in relation to
genetic diversity have been discussed by Marshall and Brown (1975), Hawkes
(1980), Chang (1985) and several other workers. For sampling of a general nature,
random collecting at predetermined intervals will be satisfactory. The intervals can
be wide in an ecologically/edaphically uniform environment or site and
proportionately small when the collecting sites include much faster changes in
topography/altitude, soil types, farming practices or such other features as observed
in an agroecologically diverse environment.
The general principle in sampling is to randomly collect a bulk sample from the
site by harvesting random fruits of a number of plants from several spots in the site.
The collector may walk across a site or a field twice – in the form of a cross or
zigzag manner avoiding sampling from the borders. This procedure of coarse grid
sampling will sample the maximum variation in the population if the site is large
and contains distinct differences in eco-edaphic conditions. In the other method,
called the clustered sampling pattern, one collects several samples within a small
area and repeats the sampling over several areas within the broad-based site. This
clustered pattern allows the inclusion of large variability related to both geographic
and microgeographic differences in environment. It may be better suited for wild or
weedy forms. In hotspot areas, the collector should collect healthy looking plants in
fields where severe disease or insect damage is evident.
Sampling Frequency and Size
The frequency of sampling (number of samples per site) and the size of each sample
would be governed much by the extent of genetic diversity and gene flow within a
taxon (cultigen or wild form) and the agroecology of the site. The collector should
use a practical approach and utilize the on-the-spot observations to devise the best
sampling technique (Hawkes 1976; Marshall and Brown 1975).
Where some populations seem to be extremely variable, one can either make
much larger samples or take several distinct samples from various parts. Non-
random collections based on ‘race identification’ can be made in addition, but
should be given distinct collection numbers and kept separate from the random
collection. Additional non-random (or selective) samples may be added as
subsamples if the collector sees any particularly interesting variant present in small
numbers, which were not included in the sample by strictly random sampling.
Collecting Sites/Sources
There are four main collecting sites that have been recognized (Hawkes1980), and
they are as follows:
1. Farmer’s fields
2. Kitchen or orchard gardens
3. Markets
4. Wild habitats
344 S. Uma et al.
However, the most important is collection from the farmers’ fields which pro-
vides visual realization of the much required wealth of landraces and primitive cul-
tivars of the main fruit crops like banana. Much would depend on the location and
size of farmers’ landholding. Subsistence farming sites are ideal for sampling rich
diversity of crops, grown in kitchen gardens/orchards for small-scale markets. The
choice would thus be site-dependent. Also, the farmers’ stores provide useful vari-
ability from the previous year’s harvest.
Collecting Wild Relatives of Crop Species
As compared to the collecting efforts made in augmenting genetic diversity in cul-
tivated plants as reflected above, relatively much less emphasis has been placed so
far on the wild relatives of crops plants. This category of diversity is required to
further enlarge the genetic base available to the breeders for crop improvement. The
wild species and the weed races represent the highest level of genetic heterozygosity
and heterogeneity among the different classes of germplasm. It is easy to exploit the
wild progenitors and other wild species in the primary and secondary gene pools
respectively, but not the tertiary gene pools.
Field Data Recording
IPGRI guidelines are followed for recording the passport data (IPGRI/INIBAP
1996) on banana.
Germplasm Collection Strategies
The general strategy adopted for germplasm collection in banana is summarized as
follows. These points would further strengthen the know-how of the collector:
1. Sucker collection should be made from each individual clump/each distinct
morpho-type in a village/hamlet.
2. Collection should be made repeatedly at regular intervals.
3. Sample should be collected at as many sites as possible according to the avail-
ability of time.
4. Choose sampling sites over as broad as environmental range as possible.
5. If considerable morphological variation is found in a population, efforts should
be made to make separate samples of each type.
6. Taking photographs of each and every part of the plant is a must; video cover-
age is also appreciable.
7. Field notes/data are very important and it should be meticulous.
8. If possible/available, seed should be also collected.
9. Labelling should be legible/clear and collection number should be tagged with
the sampling.
10. Supplement with propagules/seeds collections, where possible, and give same
collection numbers if seeds come from the same plants as the vegetative
samples. If they do not or are bulked samples, give separate collection numbers
Depletion of Musa Genetic Wealth in India

In the priority areas/regions which hold rich native genetic diversity for different
crops, the depletion of genetic resources is of different kinds. This is visible in
advanced or developed agricultural sectors where large impact of high-yielding
varieties operates as well as in other situations with ethnic diversity, agroecological
variation and botanical richness. The causes of depletion of plant genetic resources
particularly Musa genetic wealth in the North-Eastern region, Western Ghats,
Eastern Ghats and Western Himalayas are as follows:
• Spread of high-yielding varieties leads to over-exploitation of natural economic
plant wealth, etc.
• Excessive biotic interference – large-scale felling of trees/destruction of natural
vegetation, shifting cultivation, etc. There has been loss of native variability and
endemic/rare cultivated and wild types and other economic plants.
13 Establishment of the Collected Genetic Materials
In a vegetatively propagated crop like banana, suckers are used as planting materi-
als. It is very important to establish the collected suckers/propagules. Hence, imme-
diately after the collection trip, the planting materials should be thoroughly treated
with fungicide for quarantine; otherwise soil borne fungus may enter the field gene
bank. In the present scenario of conservation strategies, more specifically in banana,
in vitro conservation is also complementary for in vivo conservation, and hence it is
always advisable to have one set of samples under in vitro conservation.
14 Characterization of the Banana Genetic Resources
Simmonds and Shepherd (1955) developed simple numerical scoring system for 15
characters of banana for their classification into genomic groups below the level of
the genus. A modified and revised score card was developed by Silayoi and
Chomchalow (1987) and Singh and Uma (1996), respectively. The later one is to
encompass the genetic variability prevalent in India. The disadvantages of these
systems are the overlapping scores and their difficulty to discriminate accessions
within the same genomic and subgroups.
A general key to morphological characters of various groups was developed
based on the high degree of variability for characters like pseudostem blotching,
pigmentation, leaf length/breadth ratio and flowering – fruiting duration, number of
hands and fingers, peduncle nature, pedicel length, fruit size, taste and nature of ripe
fruit flesh, etc. in the germplasm collection – and in 1996, the Musa descriptor was
developed by INIBAP/IPGRI, which involves 121 traits for characterization. User-
friendly software was developed for classifying a banana genotype/clone by using
346 S. Uma et al.
pairwise discriminant function for five genomes accounting to 10 equations and 19

morphological characters.
Morphological traits, influenced by genotype x environmental (G x E) interac-
tions, are governed by a few or many genes. Numerical methods lack any reliable
significance test to prove their accuracy, and the whole numerical procedure must be
repeated when every time new clones were incorporated. The unreliability of some
parameters led to the large-scale adoption of molecular techniques. Complementing
morphotaxonomic characterization with molecular characterization is more useful
in drawing meaningful conclusions on diversity.
During the initial days, dominant markers like RAPDs were used for character-
ization of Musa germplasm followed by codominant markers like SSRs, but in
recent days advanced marker systems like SNPS and DArT markers are widely used
based on the expertise, infrastructure and available funding. Molecular markers
largely facilitated the genetic resource management by way of identification of
duplicates, establishment of core collection, fingerprinting, identification of genetic
contamination and quantification of genetic drift (Rao 2004). Further characteriza-
tion of the core collection using trait-specific markers enabled the establishment of
mini-cores for specific traits (Meena et al. 2010). This provides precise knowledge
on the extent of diversity in a germplasm which is a prerequisite for strategic breed-
ing in a crop like Musa that is recalcitrant for breeding owing to parthenocarpy,
sterility and polyploidy. The diversity is assessed by measurement of genetic simi-
larity (GS) or genetic distance (GD) between accessions in a genetic pool. It also
helps in tagging and mapping of agronomically important traits, marker-assisted
breeding and map-based cloning of genes. Recently diversity array technology
(DArT), which is a robust and cost-effective microarray-based technique that offers
high multiplexing, independent of sequence information (Jaccoud et al. 2001), is
being used widely for diversity analysis. ICAR-NRCB, Trichy, has analysed 186
germplasm accessions using DArT markers, and the diversity array has been eluci-
dated (Anon. 2015, 2016).
15 Conservation of the Banana Genetic Resources
The conservation of Musa genetic diversity is carried out by the following three
complementary methods:
1. In situ conservation, i.e. in the wild natural habitats where specific species
evolved and continue to do so.
2. On-farm conservation, i.e. in farmers’ fields where continuous cultivation, adap-
tation and improvement of cultivars is often carried out by small-scale farmers
cultivating traditional local cultivars (including home gardens).
3. Ex situ conservation has been defined by CBD (1992) as the conservation of
components of biological diversity outside their natural habitats, for example, in
a field research station, in vitro gene bank (medium term) and/or in
cryopreservation (long term).
Field gene bank is one of the ex situ conservation methods to keep the genetic diver-
sity as living plants either in the field or pots in a nursery. It is most suitable for
vegetatively propagated crops like banana which are recalcitrant to produce seeds
and have a long life cycle. It is often a complementary strategy to other conservation
methods such as in vitro and cryobank. The first and foremost requirement is a long-
term funding for collection, characterization and conservation under field gene
banks (Table 8).
In case of field gene bank, the merits are easy access to resources and direct
control over germplasm. They can be readily characterized and evaluated. Those
bananas stored in field gene bank have a lower risk of losing genetic integrity. This
type of conservation is most preferred for genotypes that commonly produce
variants since they could be more easily identified and rogued in the field than under
in vitro conditions. The demerits are that the field gene banks require more labour,
inputs and space for maintenance and they are highly prone to pests, diseases and
natural calamities like drought, floods, volcanoes and hurricanes, and therefore the
chances of losing the genetic material is more (Maxted et al. 1997; Engelmann and
Engels 2002; Dulloo et al. 2001). Therefore, it requires a rationalized approach
dealing with properly characterized germplasm and ensuring the management of
true-to-type germplasm. Non-adaptability to the new environment is yet another
reason for the loss of genetic material. This is frequently the case for species of
Australimusa, Callimusa and for those Eumusa species restricted to mountainous
climates.
Most of the foresaid constraints are related to agronomic problems, but in the
context of germplasm conservation in field gene bank, where we handle diverse
materials, often with little biological and ecological information, precautionary
measures should be taken to plant pest and disease-free material and to provide
utmost care and attention to establish them without any loss or damage so as to reap
its fullest potential.
Gene bank is most useful in safeguarding varieties which are nearing extinction,
not conserved in other gene banks and those which possess high economic values.
This is how many field gene banks have come into existence both at the global and
Table 8 Comparative merits and demerits of ex situ conservation methods

Parameter Field gene bank In vitro gene bank Cryobank
Requirement of technical protocols Low Medium High
Applicability for diverse germplasm accessions High Medium Low
Space requirement High Medium Low
Laboratory and infrastructure required Low High High
Technical expertise required Low Medium High
Risk to stored germplasm High Medium Low
Availability of germplasm for utilization High Medium Low
Ease in germplasm exchange Medium High Low
cost involved Medium High Low
Agrawal et al. (2007)
348 S. Uma et al.
national level. The important field gene banks are present in countries like Latin
America, Cameroon, Uganda, India, China, Vietnam, the Philippines, Indonesia,
Australia and Papua New Guinea, most of whom have contributed for the global
collections at ITC, Belgium. India has eight field gene banks located at various parts
of the country. NRCB, Trichy, and BRS, Kannara, consist of both active and base
collections, and others include mainly active collections of commercial significance
maintained exclusively for evaluation, multiplication and distribution purposes.
Field gene banks usually possess three major components. Primary gene pool – it
consists of cultivated and wild landraces. Secondary gene pool – it includes species
which can be crossed with primary gene pool and produce at least some fertility.
Tertiary gene pool – this contains species which cannot be crossed with primary
gene pool under normal conditions. But it can be facilitated through modern methods
like distant crossing, embryo rescue and chromosome doubling.
16 Establishment of the Field Gene Bank
16.1 Establishment of Virus-Free Germplasm
Indexing should be carried out primarily for four viruses, namely, Banana Bunchy
Top Virus (BBTV), Banana Streak Virus (BSV), Banana Bract Mosaic Virus
(BBrMV) and Cucumber Mosaic Virus (CMV), and should be done twice during
crop duration at 6 months stage and at fruiting. Serological tests like Enzyme Linked
Immunosorbant Assay (ELISA) (Diekmann and Putter 1996; Geering and Thomas
1996) are based on antigen-antibody reaction in which the viral coat protein acts as
antigen and specific antibody raised against coat protein serves as antibody for
immunological tests. In case of nucleic acid or viral genome-based techniques, PCR
(Dietzgen et al. 1999; Harper et al. 1999; Selvarajan et al. 2007), RT-PCR (Dietzgen
et al. 1999; Roberts et al. 2000) and Nucleic Acid Spot Hybridisation (NASH) (Mas
and Pallas 1995) have been standardized for major banana viruses and are widely
used for indexing. If the germplasm is found infected, the propagule should be
destroyed, and efforts should be made to recollect the virus-free germplasm for its
establishment in the field gene bank.
16.2 Soil and Climatic Requirements
Banana prefers a deep and well-drained loamy soils with high water-holding capac-
ity and organic matter content as reported by Purseglove (1988) and grows well in
humid tropical regions lying between 20°N and 20°S latitudes. It comes up in alti-
tudes ranging from 100 to 500 m above MSL. It requires a mean minimum tempera-
ture of above 19 °C (with the optimum average temperature being 27 °C) and
> 100 mm rainfall per month as reported by Robinson and de Villiers (2007). But
the field gene bank cannot depend on vagaries of rainfall, and hence irrigation facili-
ties should be available.
16.3 Planting System
Double the space required for the collection should be reserved for the genebank
purpose. For example, if a collection of 400 plants occupies 1 ha, 2 ha should be
made available for planting germplasm accessions. Field which has not been under
banana for the previous 2 years on it and should have been planted with a non-host
crop should be selected for the establishment of field gene bank. Soils with adequate
drainage and without any water logging problem should be selected. Normally the
site is divided into bands, which corresponds to subgroups. Planting is always
preferred in single rows with five plants per row. There should be a spacing of 3 m
between rows and 2 m within rows.
Best planting materials are sword or maiden suckers which do not possess broad
leaves until they are more than 1 m high. Planting material should be selected at the
time of flowering or at the end of the growing season. They should be free from
undesirable variations and other pests and diseases.
16.4 Ensuring Accession Identity
Accession identity is monitored by comparing the plant appearance, male bud and
bunch characteristics with the passport data. For these purposes, Musa descriptor,
Musalogue and Musa Germplasm Information system (MGIS) could be used. If the
traits match with those of reference, then it is true to type (TTT). If not, then it is
either mislabelled (ML) or off type (OT). If ML, true identity is sought, and if OT,
it is destroyed, and efforts should be made to reintroduce the accession, for
monitoring the accession identity.
17 Problem in Maintenance of Field Gene Banks
17.1 Mislabelling and Mix-Ups
Proper labelling should be done by a well-trained staff because they are primarily
responsible to register and assign sequential numbers. Horticulture and botanical
taxonomy should be validated once in every 3–5 years. Large land area adds to the
cost of maintenance, so duplicate accessions may be identified with a crop specialist
350 S. Uma et al.
or using markers, and they should be eliminated. Secondly it is better to maintain

fewer duplicates for those accessions which are quite familiar and those which are
maintained/available at other gene banks.
17.2 Loss of Diverse or Inaccessible Genes
There is every chance of losing diverse or inaccessible genes during the process of
elimination of duplicates. No accession should be eliminated based on the evaluation
of a single trait. Disease indexing methods are quite expensive; hence it is neglected/
ignored many a times leading to inadvertent disease spread. Hence, attempts should
be made to develop/design methods that are less laborious and time-consuming like
those of dipstick technique available for detection of banana viruses.
To curtail the pest and disease spread and for easy maintenance, it is always bet-
ter to plant the resistant and susceptible ones separately. Similar caution should be
taken in case of salt-tolerant and susceptible varieties taking care to plant the sus-
ceptible ones in salt-free areas. AA diploid are slender with short duration and
require only less fertilizers. Contrarily BBB are robust with longer duration and
require more amount of fertilizers. So, it is again better to plant such accessions
separately. Knowledge on soilborne diseases is essential to decide an appropriate
site for germplasm.
17.3 Slow Rate of Multiplication
It is mainly due to hormone-mediated apical dominance. Non-availability of ade-

quate planting materials in a shorter time frame often delays the MLT evaluation.
Hence, we need to adopt macro-propagation techniques for rapid multiplication.
Most often the suckers of wild germplasm collected from various parts of India fail
to establish in the field gene bank because of the extreme weather conditions
prevailing at the site of establishment. Further the suckers of wild germplasm are
huge and heavier posing difficulty in transportation. Under such circumstances, it is
easier to carry fruits which are profusely seeded and could be germinated either
under ex vitro and in vitro conditions. Ex vitro germination is usually done in sand
beds where the germination is highly erratic and poor (< 0.01 per cent). But the
germination per cent is greatly improved by hydro or hormonal priming of seeds
followed by excision and culturing of embryos on MS medium supplemented with
or without growth hormones (Arun et al. 2013).
18 In Vitro Conservation
It is essential to adopt alternative strategies for those accessions which are unique
and nearing extinction. Duplication of field gene banks in more than one site is
mandatory and ideally in an in vitro gene bank as a safety backup. Fewer duplicates
are sufficient for those accessions available at other gene banks except for unique
ones. Gene bank site should have adequate rainfall or water supply for supplementary
irrigation. It should be in a secured site free from theft and other encroachments.
Newly introduced collections should be closely observed for the first one or two
crop cycles. Once after receipt of the material, they should be thoroughly disinfected
by an entomologist and pathologist. If imported, ICAR-NBPGR, New Delhi, will
take care of this, while indigenous collection should be screened for inherent
diseases like virus. Virus indexing is a must. Virus-free accessions should be
maintained either in vitro or under special enclosures.
To bring down the cost and minimize the loss of germplasm accessions, research
should be emphasized on the following areas:
1 . Development of low-input maintenance strategies
2. Development of optimal screening procedures to avoid the pest- and disease-
affected material during collection and their introduction in field gene banks
3. To study and understand the specific environmental requirements of different
banana accessions in order to better manage them in field gene bank
18.1 In Vitro Gene Bank
Biodiversity hotspots around the globe are at risk, and in vitro propagation methods
have been used for rescuing and conserving endangered plants in many countries. In
vitro propagation and conservation also contribute to the maintenance of natural
populations through the reintroduction of preserved material to the original habitat
(Pence 2011). In vitro gene banks are easily established in a crop like banana which
already has a well-developed tissue culture system. The accessions prioritized for
conservation under in vitro gene banks are those that (1) have rare genes or gene
combinations, (2) are difficult to maintain in the field, (3) are not readily available
at the active gene bank sites and/or (4) are exotic and have been introduced with
great efforts (Reed et al. 2011). The most important constraint under in vitro
conservation of plant materials is the cost involved in their maintenance in terms of
time, manpower, infrastructure, power consumption, etc. and frequent occurrence
of contamination including endogenous bacterial and fungal infection. Exudation of
polyphenols is the major problem hindering the multiplication of some banana
accessions leading to their low survival rates. The frequent occurrence of negative
somaclonal variants often poses a problem in the in vitro conservation of germplasm
(Côte et al. 1993; Vuylsteke and Ortiz 1996; Rodriguez et al. 1998). Although the
origin of off-type plants is not understood, chromosome number changes (polyploidy,
352 S. Uma et al.
aneuploidy) due to repeated in vitro subcultures might have contributed for this
phenomenon. More research is needed to standardize protocols for accessions
which are recalcitrant for in vitro tissue culture multiplication.
18.2 DNA Bank
It is possible to store the DNA in three different forms, namely, total genomic DNA,
DNA libraries and cloned DNA fragments. It is the most convenient experimental
material, easy to exchange and ready for further manipulations. The demerit is that
it allows only handling of single genes but not the whole genome. ITC established
a lyophilized tissue collection in 2004 that holds leaf samples of 883 accessions (as
on 2014). It is also a cost-effective way to preserve the molecular materials and
representative information from each species and cultivar in the ITC, serving as a
future reference for the identification of accessions in the active and base collections.
Similarly, ICAR-NRCB, Trichy, has established a DNA bank as early as 2008 which
conserves the DNA of the core collection accessions (360 nos.) in a deep freezer at
−80 °C. They are withdrawn at regular intervals of 4 years and checked for their
stability, and fresh extractions are made for those germplasm, whose DNA has
degraded and replenished to the bank (Anon. 2011). This would be of immense use
for population genetics analysis. However, the aim is to make a wide range of
diversity readily available at low cost to molecular scientists who are using DNA for
their studies. Future research should focus on the conservation of DNA libraries and
cloned DNA fragments.
18.3 Cryoconservation
Cryopreservation is becoming an increasingly used method for the long-term stor-

age of plant genetic resources (PGRs). Cryopreservation requires only a minimum
of space and low level of maintenance.
18.3.1 Short- and Medium-Term Conservation
Slow-growth strategies are used in a number of institutions throughout the world for
the preservation and distribution of clonally propagated plant germplasm (Ashmore
1997). It is a versatile tool that can provide access to disease-indexed plants for
distribution to scientists or farmers across the globe. Medium-term conservation is
obtained under slow-growth conditions by considering (1) physiological stage of
the explant, (2) osmotic agents and growth inhibitors, (3) reduced temperature,
(4) medium alterations such as reduced mineral or sucrose concentrations, (5)
reduced oxygen or (6) alginate encapsulation (Zee and Munekata, 1992; Dulloo
et al. 1998; Engelmann 1997, 1999; Harding et al. 1997; Oka and Niino 1997; Reed
et al. 2005; Rai et al. 2008; Sarasan 2011). Minimal growth method is desirable for
in vitro germplasm conservation as it reduces the subculture frequency resulting in

minimal occurrence of somaclonal variations. Selection of explant is also important
for in vitro conservation as cultured cells and callus that may occur are more prone
to somaclonal variations compared to in vitro shoot cultures. Hence, there is a
preference for using shoots for in vitro conservation. Meristem culture under limited
growth conditions is being currently applied for ITC collection at KUL, Belgium.
This is considered suitable for ‘active’ collections of banana germplasm, and this
could be readily used for germplasm exchange.
18.3.2 Long-Term Conservation
Cryopreservation is based on the reduction and subsequent interruption of meta-

bolic functions of biological materials by decreasing the temperature with LN2
(−196 °C) while maintaining viability. At −196 °C, almost all the cellular metabolic
activities are quiescent, and the cells can be preserved in for a long term. It is essen-
tial to avoid lethal intracellular freezing that occurs during rapid cooling in LN and
warming in order to maintain the viability of hydrated cells and tissues (Sakai and
Yoshida 1967). Cells and tissues that are to be cryopreserved in LN2 need to be
sufficiently dehydrated before being immersed in LN2. These new techniques have
facilitated the cryobanking of other plant species that have established tissue culture
techniques like Musa germplasm. The cost is much lower than for active storage.
Experiments have been carried out with seed, zygotic embryos, somatic embryos
and embryogenic cell suspensions. It is not feasible to apply these approaches to all
varieties and cultivars since, firstly, only a limited number of them produce seed of
which the genetic make-up is unknown. Secondly, only a few produce somatic
embryos and embryogenic cell suspensions even after months of in vitro culturing.
The cryopreservation of readily available banana meristems has therefore been
considered, despite their apparent recalcitrance to existing techniques. Currently,
three cryopreservation protocols are available for shoot meristematic tissues of
banana: (1) simple freezing of proliferating meristem cultures using a sucrose
preculture (Panis et al. 1996), (2) vitrification of apical meristems (Thinh et al.
1999) and (3) vitrification of sucrose-precultured meristem cultures (Panis et al.
2000). Simple cryopreservation method could be used for long-term storage of
banana ‘base’ collections under more secure, low-input conditions. Slow freezing
and use of highly concentrated vitrifying solutions were both totally ineffective,
while the encapsulation-dehydration method developed by Fabre and Derreudre
(1990) resulted in a low survival rate of 8.1% for banana (Panis and Swennen 1994;
Panis 1995). The encapsulation-dehydration method is labour-intensive and time-
consuming since it involves many steps and excision of small (1 mm)-sized
meristems. Since banana shoot tips have proven to be extremely sensitive to
dehydration and unresponsive to slow freezing (Panis 1995), methods involving
rapid freezing without an additional dehydration phase are essential.
Very recently it has been shown that cryopreservation could also be used for
eradication of viruses (Helliot et al. 2001). Cryotherapy-based procedures are easy
to implement and do not require special equipment in addition to those typically
354 S. Uma et al.
available in a plant tissue culture laboratory. Cryotherapy facilitates treatment of

large numbers of samples. It yields pathogen-free plants at a high frequency avoiding
the difficulties associated with the ecision of very small meristems.
So far International Transit Centre (ITC), Belgium has cryopreserved 938
accessions. A replicate set of cryopreserved material is held at the IRD in
Montpellier, France, with 801 accessions to date. This provides an off-site backup
as a further safety measure under the ‘black-box’ arrangement wherein the mate-
rial cannot be touched without requesting the depositor and returned on request.
ICAR-NBPGR, New Delhi, has successfully cryopreserved 92 banana germ-
plasm in the form of proliferating meristems and found that neither cryopreserva-
tion nor time lag has affected the genetic stability of the cryopreserved germplasm
(Anon. 2011). Attempts are in progress to cryopreserve various explants of banana
including seed, embryo and pollen. If cryopreservation of ECS is successful, then
ECS could be maintained at a minimal cost in a centralized facility from where it
could be distributed to various production facilities depending on their requirement
for further regeneration of quality planting material. Poor viability of banana seeds
necessitates the development of protocol for cryoconservation of seeds. Likewise,
development of protocol for storage of pollen is a must in banana since the flowering
in male and female parent often does not synchronize for effecting pollination
towards the successful development of banana hybrids. By using the cryoconservation
technique for pollen storage, the need for maintenance of male parents could be
eliminated, and it could be easily exchanged across breeding centres.
19 U
tilization of Banana Genetic Resources in Crop
Improvement
Characterization and evaluation of banana genetic resources are essentially required

to study, understand and enhance the conservation and their utilization in crop
improvement. The use of crop wild relatives (CWR) genes in crop improvement is
a well-established fact (Hajjar and Hodgkin, 2007). Though banana is recalcitrant
for breeding, the cultivated diploids and wild Musa species are frequently used for
their genetic improvement. For example, Calcutta 4′ (Musa acuminata ssp.
burmaniccoides), a wild, nonedible diploid banana, is used to impart resistance to
black Sigatoka, the most serious constraint to banana production globally caused by
the fungus Mycosphaerella fijiensis (Carlier et al. 2002). The progenies were found
to be resistant to Sigatoka and fusarium wilt. But often, the hybrids inherit only the
resistant traits deteriorating the original quality of the cultivar to be improved
(Saraswathi et al. 2016). Therefore, in recent past, molecular breeding is being
given impetus, and it is in progress at a greater pace which has resulted in the
successful identification and isolation of full-length genes conferring resistance to
biotic and abiotic stresses. Pectin methyl esterase inhibiting genes have been
isolated from banana cv. Harichal which is likely to delay ripening (Srivastava and
Dwivedi 2000) and PSY genes responsible for synthesis of provitamin A from cv.
Asupina (Mlalazi et al. 2012). Likewise, RGAs conferring nematode (Backiyarani

et al. 2013) and fusarium wilt resistance (race 4) (Usharani et al. 2016) have been
isolated from Musa spp. Incorporation of such desirable genes into banana through
cis-/transgenic approaches enables the sustainable utilization of genetic resources in
banana improvement.
20 Conclusion and Recommendations
Banana with its limited diversity is difficult to be bred conventionally owing to the
parthenocarpic, sterile and polyploidy nature of the crop. However, serious attempts
have to be made to tap the potential of available diversity for the improvement of
banana. This necessitates the strategic collection of genetic resources available
across the globe through exploration. The collections have to be systematically
conserved for subsequent characterization, evaluation and utilization. The
collections should be necessarily indexed for major four banana viruses using
sensitive techniques prior to its establishment in the field gene bank. This will avoid
the inadvertent spread of viruses which may wipe out the entire germplasm if left
unattended. In recent days, in situ conservation is being given impetus under the
watch and ward of ethnic tribal groups and non-governmental organizations. Careful
planning and field management helps to mitigate the problems encountered in the
field gene banks. Periodically the germplasm has to be monitored for genetic fidelity
to maintain the true to type as the crop is highly prone for natural mutations.
Rejuvenation of wild Musa species is usually taken up using selfed progenies.
It is always better to develop a complementary conservation strategy each appro-
priate to a specific component part of the overall conservation programme and taken
together in such a way that they complement each other for the most efficient and
safest conservation in the long term. GIS could also be applied to the genetic
resources conservation as it has become a reliable tool to visualize and analyse spatial
patterns in genetic data in relation to ecological data. Complete characterization
should be accomplished through morphotaxonomic and molecular means so as to
have knowledge even on the minute details of the valuable resources inclusive of
their phylogenetic relationships that might enhance the rate of success in our
breeding endeavours. The job of a gene bank curator does not end with
characterization of collected resources; it has to be carefully documented for the
benefit of the scientific community to study their evolutionary pattern in future. The
characterized germplasm should be evaluated for their response to various biotic
and abiotic stresses and used appropriately in the improvement of already existing
cultivars. The gene bank curators should remain as unsung heroes over long-term
conservation programmes of crop plants.
356 S. Uma et al.
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Banana Genetic Resources

S. Uma, M. S. Saraswathi, and P. Durai

Keywords Banana · Genetic resources · Collection · Characterization ·

1 History, Origin and Evolution

S. Uma (*) · M. S. Saraswathi · P. Durai

© Springer Nature Singapore Pte Ltd. 2019 321

M. textilis, a member of Australimusa, is endemic to the Philippines. Several

Banana is a large, herbaceous member of the family Musaceae belonging to the

4 Phylogeny, Domestication and Dispersal

5 Global Banana Uses, Production and Trade

Table 1 Major banana-producing countries of the world

average production of 15–20 suckers/year/clump. They have attractive flower buds

5.1 Export Scenario of India

Fig. 1 Banana export by different countries during 2017

Fig. 2 Major Indian banana-importing countries

Table 2 Major banana varieties distributed across the globe

in Thailand, Choui in Vietnam, Kela/Vazhai/Vazha/Bale and Arati in India and

6 Status of Wild Musa in India and Their Distribution

Table 3 Distribution of wild Musa species in India

Table 4 Distribution of recently identified Musa species in India

Table 5 Distribution of Rhodochlamys and allied genus Ensete in India

parthenocarpy which was later confirmed by RFLP studies (Carreel 1994).

8 Polyploid Bananas (Triploids and Tetraploids)

8.1 Bispecific Polyploids

Introgression of acuminata diploids with M. balbisiana occurred either in their pri-

8.2 AAB Cultivars

Introgression between two species in various combinations, especially M. acumi-

et al. (1993) with M. acuminata ssp. banksii contributing to AA genomes in both

8.3 ABB Cultivars

9 Status of Cultivated Bananas in India

10 Evolution of Ensete

Table 6 Major banana-growing states and the common cultivars

11 Exploration and Collection of Musa Genetic Resources

12 Technical Guidelines for Exploration

Information on the locations proposed to be visited, routes to be followed, position

12.1 Logistic Preparation/Implementation

Table 7 Important items and equipments required

12.2 Practical Guidelines

12.2.1 Survey Tactics

germplasm in sites/villages much away from the approachable roads, as the

12.3 Knowledge on Agroecology and Musa Distribution

It is necessary to acquire knowledge on the agroclimatic conditions in relation to the

12.4 In Situ Observations

2 . Minimum description of the species/landrace/variety

12.5 Germplasm Collection

especially in physiological characters. Sampling methods used must ensure the

Depletion of Musa Genetic Wealth in India

13 Establishment of the Collected Genetic Materials

14 Characterization of the Banana Genetic Resources

pairwise discriminant function for five genomes accounting to 10 equations and 19

15 Conservation of the Banana Genetic Resources

Table 8 Comparative merits and demerits of ex situ conservation methods

16 Establishment of the Field Gene Bank

16.1 Establishment of Virus-Free Germplasm

16.2 Soil and Climatic Requirements

16.3 Planting System

16.4 Ensuring Accession Identity

17 Problem in Maintenance of Field Gene Banks

17.1 Mislabelling and Mix-Ups

1 History, Origin and Evolution

4 Phylogeny, Domestication and Dispersal

5 Global Banana Uses, Production and Trade

5.1 Export Scenario of India

6 Status of Wild Musa in India and Their Distribution

8 Polyploid Bananas (Triploids and Tetraploids)

8.1 Bispecific Polyploids

8.2 AAB Cultivars

8.3 ABB Cultivars

9 Status of Cultivated Bananas in India

10 Evolution of Ensete

11 Exploration and Collection of Musa Genetic Resources

12 Technical Guidelines for Exploration

12.1 Logistic Preparation/Implementation

12.2 Practical Guidelines

12.2.1 Survey Tactics

12.3 Knowledge on Agroecology and Musa Distribution

12.4 In Situ Observations

12.5 Germplasm Collection

13 Establishment of the Collected Genetic Materials

14 Characterization of the Banana Genetic Resources

15 Conservation of the Banana Genetic Resources

16 Establishment of the Field Gene Bank

16.1 Establishment of Virus-Free Germplasm

16.2 Soil and Climatic Requirements

16.3 Planting System

16.4 Ensuring Accession Identity

17 Problem in Maintenance of Field Gene Banks

17.1 Mislabelling and Mix-Ups

17.2 Loss of Diverse or Inaccessible Genes

17.3 Slow Rate of Multiplication

18 In Vitro Conservation

18.1 In Vitro Gene Bank

18.2 DNA Bank

18.3.1 Short- and Medium-Term Conservation

18.3.2 Long-Term Conservation

20 Conclusion and Recommendations