Overview of Banana and Plantain (Musa spp.

) Improvement in Africa:
Past and Future
J. Lorenzen1, A. Tenkouano2, R. Bandyopadhyay2, B. Vroh2, D. Coyne1 and L. Tripathi1
International Institute of Tropical Agriculture, P.O. Box 7878, Kampala, Uganda
2
International Institute of Tropical Agriculture, PMB 5320, Ibadan, Nigeria

Keywords: black leaf streak, breeding, Fusarium wilt, nematodes, resistance, weevils
Abstract
Since an unrecorded introduction from Asia in prehistoric times, banana and
plantain (Musa spp.), commonly called bananas, have become major food and cash
crops in Africa. The 4 million ha of bananas in Africa represent nearly a third of
global production. Increased movement of plant material in the past century also
introduced pests and diseases that became new constraints to banana production in
Africa and have destabilized banana production with susceptible traditional
landraces. Biotic challenges to banana production in the region include fungi,
bacteria, viruses, nematodes and insects. Introducing host-plant resistance, whether
by conventional breeding or by biotechnology, is the most economical and
sustainable means of managing pests and diseases. In recent decades, progress has
been made in identifying sources of host-plant resistance, identifying germplasm in
other countries most like original progenitors of African landraces and developing
genomic tools to increase the efficiency of developing resistant lines. Major
programs for banana breeding in Africa are located in Nigeria (International
Institute of Tropical Agriculture), Cameroon (Centre Africain de Recherche sur
Bananiers et Plantains), and Uganda (National Agricultural Research
Organization/International Institute of Tropical Agriculture). Banana breeding is
slow and land intense compared to annual crops, so increasing breeding efficiency is
a valuable objective. Good progress has been made in introgression resistance to
black leaf streak disease, burrowing nematodes, banana weevils, and Fusarium in
elite selections. There is room for progress in producing high-yielding cultivars,
resistant to multiple biotic threats with similar organoleptic qualities as traditional
cultivars.
INTRODUCTION
Banana and plantain (Musa spp.), commonly called banana, is an important world
food crop that is far more important to food security and livelihoods of millions of
smallholders in tropical countries of Africa, Asia and South/Central America than its
western image of an industrial export fruit would suggest. The vast majority of global
Musa production is for domestic consumption, with a high proportion of production
coming from small farms and gardens. From early domestication in Southeast Asia and
the islands extending toward Australia, banana spread to Africa in undocumented
introductions, long before recorded history for that region. Phytolith evidence suggests
that banana reached Africa several millennia ago, with relatively rapid penetration into
Central Africa (Mbida et al., 2001; Lejju et al., 2006). Although we don't know whether
banana reached Africa as already cultivated triploids or as diploid precursors, the very
limited genetic diversity within the major classes of African plantains (AAB genome) and
East African highland bananas (EAHB, AAA genome) suggests that it may likely have
been triploid cultivars that spread inland from the coast(s). The relatively high genetic
diversity of Musa in islands off the East African coast (Simmonds, 1966) and the
mountains of Tanzania may be remnants of earlier introductions or have come in later
waves of Indian Ocean trade.
BANANA BREEDING PROGRAMS IN AFRICA
The Musa breeding programs in Africa were relatively late to start and could build
Proc. IC on Banana & Plantain in Africa
Eds.: T. Dubois et al.
Acta Hort. 879, ISHS 2010

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on the theoretical and practical foundations of earlier breeding efforts (see below).
Although the Musa research program at the International Institute of Tropical Agriculture
(IITA) was initiated at its high rainfall station in Onne, Nigeria in the mid-1970s, the
breeding effort did not commence until the imminent threat of Black leaf streak virus
(BLSV, caused by Mycosphaerella fijiensis) led several countries in the region to request
IITAs assistance in breeding for this serious foliar pathogen (Vuylsteke et al., 1993a, b,
1997; Ortiz, 2001). Another regional effort, the Centre de Recherches Rgionales sur
Bananiers et Plantains (CARBAP) was initiated in the late 1980s with the assistance of
the French research and development agency, Centre de Coopration International en
Recherche Agronomique pour le Dveloppement (CIRAD) (Jenny et al., 1994). In
response to the dispersal of BLS across East Africa, and in light of the very high
importance of cooking bananas to food security in that region, the East African banana
breeding program was initiated in 1994 as a cooperative venture between the East and
Southern Africa Regional Center of IITA and the National Agricultural Research
Organization (NARO) of Uganda (Ortiz, 2001).
IITAs activities in Musa breeding at Onne focused on recovering plantain traits
by crossing fertile plantain cultivars with diploids with desired resistance to BLSV and
other diseases, and generating resistant secondary triploids by crossing tetraploid hybrids
with improved diploids. Although early selections were mostly tetraploid (e.g., Vuylsteke
et al., 1993c, 1995), those tended to be unpredictable with regard to seed set and the
program moved to develop secondary triploids.
RESISTANCE BREEDING
Fusarium oxysporum
Fusarium wilt (syn. Panama Disease, caused by Fusarium oxysporum f. sp.
cubense) was the driving force for initial attempts to breed bananas by both public
(British, Trinidad) and private efforts (Buddenhagen, 1990). Fusarium wilt has had a huge
influence on the banana industry and was described in early reports according to races
(14), defined by differential genotypes (Stover and Simmonds, 1987). More recently, the
complexity of Fusarium banana pathogens have been described using vegetative
compatibility groups, of which there are at least 25 (Ploetz and Pegg, 2000). Resistance to
Fusarium wilt was an essential and integral part of the very large private breeding effort
that was later donated to the Fundacin Hondurea de Investigacin Agrcola (FHIA) in
Honduras (Rowe and Rosales, 1996). Although this effort largely focused on races 1 and
2, an improved diploid, SH-3362 (AA genome), also showed resistance to tropical race
4 (Rowe and Rosales, 1996). Resistance to Fusarium tropical race 4 has also been
reported in segregating populations derived from intercrossing wild accessions of Musa
acuminata subsp. malaccensis (AA genome) (Javed et al., 2004). With the spread of this
aggressive race in Asia, and fears that it may affect the banana industry in the Americas
or Africa, more urgency has been placed on finding and deploying sources of resistance to
these strains.
Black Leaf Streak Virus
BLSV is a highly destructive foliar disease of banana (Pasberg-Gauhl et al., 2000;
Marn et al., 2003). It likely originated in the West Pacific isles, Australasia and/or the
Southeast Asia region (Pasberg-Gauhl et al., 2000). BLSV causes much higher levels of
damage and decreased yields than the Yellow Sigatoka (caused by Mycospaerella
musicola leach) pathogen that spread globally much earlier (Stover, 1962; Jones, 2000). It
arrived in Africa in the early 1970s, about the same time that it was reported in Latin
America (Pasberg-Gauhl et al., 2000). In warmer tropical areas, it has tended to displace
M. musicola (Jones, 2000).
Host plant resistance to BLSV (e.g., Four et al., 2000) provided the basis of the
original IITA Musa breeding programs in both West and East Africa (Swennen and
Vuylsteke, 1990; Vuylsteke et al., 1995). One of the most widely used sources has been
596

Calcutta 4 (AA genome), which has been a highly fertile pollen source and has yielded
resistant progeny (Swennen and Vuylsteke, 1990; Ortiz and Vuylsteke, 1994; Tenkouano
et al., 2003). Resistance in Calcutta 4 was described as being controlled by a three gene
model for partial resistance (Ortiz and Vuylsteke, 1994), although others have described
Calcutta 4 as showing a typical hypersensitive response (HR) (Four et al., 2000). In
general, a disease-slowing model of partial resistance would be more likely to be durable
than a gene-for-gene hypersensitive reaction (Flor, 1956, 1971; van der Planck, 1982).
The more desirable partial resistance associated with slower development of disease
lesions has been observed in field trials in sites with high disease pressure (Fullerton and
Olsen, 1995; Four et al., 2000; Carlier et al., 2003). The HR response to BLSV is
reported to occur in several of Musa acuminata subspecies, including banksii,
burmannica, malaccensis, microcarpa, siamea and truncata (AA genome) (Four
et al., 2000). Other resistant diploid (AA) genotypes used in breeding lines include
malaccensis accession, Pisang Lilin (HR response) and microcarpa accession, Tjau
Lagada (slow lesion development) (Four et al., 2000; Tenkouano et al., 2003). Two
IITA resistant improved diploid parents, TMB2x5105-1 and TMB2x9128-3 (AA
genome), resulted from crosses with these resistance sources (Tenkouano et al., 2003).
The concern with gene-for-gene resistance is that it will not be durable (van der
Planck, 1982). High genetic diversity among M. fijiensis populations has been reported
from near the center of origin, while relatively high diversity is also found among isolates
from other continents (Carlier et al., 2003; Zandjanakou-Tachin et al., 2009). Among
diverse isolates from Pacific islands, some overcame the resistance in genotypes
characterized as resistant, including Calcutta 4, Yangambi Km5 (AAA genome),
Paka (AA genome) and newly developed resistant hybrids (Fullerton and Olsen, 1995;
Craenen and Ortiz, 2003). Similarly, the resistant hybrid Goldfinger (syn. FHIA-01
and SH-3481, AAAB genome) succumbed to BLSV shortly after being introduced to
Samoa (Daniells, 2009). Such diversity in virulence near the center of origin suggests that
breeders should be cautious in using HR-type sources of resistance with a long-term
objective of breeding durable resistance.
Plant Parasitic Nematodes
The burrowing nematode (Radopholus similes) is one of the most destructive
nematode pests of banana. It appears to have arrived in East Africa in the 1960s, where it
has become widespread and a major limitation to banana production (Price, 2006). One
source of resistance to R. similis is AA cultivars of the subgroup Pisang Jari Buaya
(PJB) (Wehunt et al., 1978; Quneherv et al., 2009). This resistance was incorporated
into the widely-used diploid parent from the FHIA program, SH-3142 (AA genome)
(Pinochet and Rowe, 1979) and its progeny SH-3362. Yangambi Km5 (Ibota type)
has demonstrated resistance to R. similis (Wehunt et al., 1978; Pinochet and Rowe, 1979;
Fogain and Gowen, 1998) and Pratylenchus goodeyi (Fogain and Gowen, 1998), although
strains of R. similis in East Africa have overcome this resistance (Dochez, 2004).
Yangambi Km5 has not been successfully used in breeding. Unfortunately, tested lines of
the PJB subgroup tended to be susceptible to P. coffeae, although some variability among
cultivars was noted (Quneherv et al., 2009). Multiple resistances are highly desired in
breeding parents. Two diploid parents mentioned above for BLSV resistance,
TMB2x5105-1 and TMB2x9128, are also resistant to R. similis (Tenkouano et al.,
2003).
Diversity among R. similis isolates with regard to pathogenicity has been observed
to exist. Some Uganda isolates that may have originated in Sri Lanka (Price, 2006) were
more aggressive with respect to pathogenicity on banana (Plowright, 2000; Dochez,
2004). A Uganda isolate was more virulent on banana than one from Indonesia (Elsen et
al., 2004). Several authors noted PJB was susceptible to one or more R. similis isolates
from Uganda (Plowright, 2000; Dochez, 2004; Elsen et al., 2004). Yangambi Km5 was
resistant to an aggressive isolate from Western Uganda that overcame the resistance of
PJB but was not resistant to an aggressive isolate from Central Uganda (Plowright, 2000).
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A relatively rapid single root screening assay has been developed through inoculation of
single roots (De Schutter et al., 2001; Dochez et al., 2005). This assay was used to screen
a segregating diploid population for resistance to R. similis (Dochez et al., 2009) and has
since been developed for multiple nematode species screening (Coyne and Tenkouano,
2005).
In West Africa, R. similis has traditionally been the key nematode pest, and hence,
one important focus for Musa breeding programs (Tenkouano et al., 2003; Tenkouano
and Swennen, 2004). However, in recent decades P. coffeae or Pratylenchus spp. have
become increasingly prevalent and damaging and are now the primary nematode problem
in some countries and areas (Coyne, 2009). Consequently, the focus within breeding
programs needs to take account of this (Lorenzen et al., 2009), as sources of resistance
against R. similis do not necessarily confer resistance against Pratylenchus spp. (De
Waele and Elsen, 2002; Quneherv et al., 2009).
Banana Weevil
The banana weevil (Cosmopolites sordidus Germar) is the most destructive insect
pest of banana. It is thought to have originated in the Indo-Malayan region (Simmonds,
1966). This important pest has spread around the world. Host plant resistance to this
corm-burrowing pest is available, including diploids such as Calcutta 4, Kisubi (AB
genome), TMB28075-7, TMB27197-2 and TMB26142-1 (AA genome)
(Kiggundu et al., 2003), East African AA diploids Njeru and Muraru (Musabyimana et
al., 2000), as well as in Yangambi Km5 and Musa balbisiana (BB genome) (Fogain and
Price, 1994). Host plant resistance to banana weevil has been reported in AA diploids
such as Calcutta 4 and Pisang Lilin (Ortiz et al., 1995).
RESISTANCE SCREENING
A tool that is very useful for accelerating breeding is a rapid assay that allows the
differentiation between resistant and susceptible genotypes in a reasonably short period of
time. A general summary of availability of such tools for specific pathogens is given in
Table 1.
Breeding Process and Achievements
The breeding process used at IITA has been similar to other breeding programs
(e.g., Rowe, 1984, 1987; Shepherd, 1994; Tomekpe et al., 1995) but with a focus on the
target market classes of plantain and EAHB. In each case, a primary objective is to
generate resistant hybrids with similar fruit properties as the original susceptible preferred
local cultivars. This scheme has had three main components, the main one being to
produce primary tetraploids through crosses with resistant diploids and then use those as
females to produce secondary triploids. The second is to use recombination from other
sources to try to recreate the target ideotype. The third is to work on improvement of the
AA diploids (Swennen and Vuylsteke, 1990). Both banana and plantain types suffer from
the low germination rates experienced by other banana programs, necessitating embryo
rescue as the primary means to germinate seeds from most crosses. Since Musa shows
plasticity with regard to ploidy of both male and female gametes, it is also necessary to
check ploidy of all progeny that are produced. While main cultivars in all market classes
lack resistance to BLSV, only some classes lack resistance to Fusarium oxysporum f. sp.
cubense. In vitro seedlings are multiplied, before transferring them to a humidity chamber
for rooting and subsequent hardening in a screenhouse prior to field planting of duplicate
plants (retaining backup plants for gap filling). Plant development characteristics (height,
suckering, time to flower, time to fill, bunch traits), disease scores, yield and organoleptic
quality are evaluated for several fruiting generations before advancing promising
selections to advanced trials. Several promising plantain hybrids have emerged from this
program in West Africa, as well as improved diploids (Tenkouano and Swennen, 2004).
In the collaboration between NARO and IITA in Uganda, the early focus was to
identify triploid cultivars with male or female fertility that could be used to produce
598

tetraploid parents with an EAHB genetic background (Ssebuliba et al., 2006, 2008).
Improved diploids from FHIA or the IITA Onne program were generally imported.
Hybrid seedlings are produced by both programs and shared as appropriate. This
collaborative effort between IITA and NARO has resulted in some very encouraging lines
that are now being multiplied and tested more widely with farmers. Positive traits
incorporated in these secondary triploid hybrids include resistance to BLSV, Fusarium
race 1, R. similis and the banana weevil. In a field trial in four central/western districts
with three advanced hybrids and one local check cultivar, the hybrids proved much more
resistant than the local check, Mbwazirume (AAA genome), to BLSV with yield
increases of 2042%. While the eating quality of many hybrids has often been rated much
lower than the local check with regard to color (yellow preferred), texture (soft preferred)
and flavor, a few of the most advanced, high-yielding hybrids have acceptable eating
quality. In places where the combined effects of BLSV, R. similis and Xanthomonas wilt
have seriously impacted banana longevity and productivity, high-yielding hybrids with
acceptable quality may be adequate. Some recently selected hybrids have quality scores
nearly equivalent to local cultivars, indicating the future promise for successful
replacement cultivars.
Similarly, highly promising secondary triploid hybrids have been identified in the
plantain improvement program, such as PITA-14 (AAAB genome) and more recent
selections (Tenkouano and Swennen, 2004).
CONCLUSION
In answer to the question posed by the organizers of this symposium Is
conventional breeding obsolete?, the answer would be a resounding No, not at all!
Early resistance breeding has proved highly promising with regard to both plantain and
East African type hybrids, with every reason to expect that later selections will be
improved with regard to quality attributes. Some lines have resistance to multiple biotic
threats (BLSV, Fusarium, weevils, nematodes), although we currently lack suitable
resistance sources for two serious pathogenic threats (banana Xanthomonas wilt and
Banana bunchy top virus). It is becoming clear that we need to broaden the base of
resistance to important pathogens by seeking more resistance sources, and there is a need
for investigating the scientific basis for important traits like food quality and drought
tolerance. While biotechnology will develop many useful tools, the narrow genetic basis
of most market classes of bananas is a threat to the industry that can best be met by
classical breeding.
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Tables

Table 1. Availability of resistance sources and high throughput screening assays for
efficient breeding for listed biotic constraints.
Problem
Black Sigatoka
Fusarium wilt
Banana weevil
Burrowing nematode
Other nematodes
Banana Xanthomonas wilt
Banana bunchy top virus
1

Availability of
High throughput
Resistance?
screen?
Y
Y1
Y
Y2
Y
(Y)3
Y
(Y)4
Y
(Y)4
N
(Y)5
N
NNA

Donzelli and Churchill, 2007; Twizeymana et al., 2007.

Javed et al., 2004.
3
Kiggundu et al., 2007.
4
De Schutter et al., 2001; Dochez et al., 2005.
5
Tripathi et al., 2008.
2

603

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