PROMUSANews

2nd Global meeting INIBAP is presently organising the second global PROMUSA meeting. This meeting is due to be held in Cameroon in Novem- ber 1998 and will be held in association with a meeting of the Principal Inves- PROMUSA tigators of the World Bank/FAO/CFC Improvement Project, just before the International Symposium on A Global Programme for Improvement and Food Security. Two days (tentatively November 8 - 9) have been set aside for the PROMUSA meeting. It is anticipated that separate meetings of most of the A banana breeder’s response to “The Global working groups of PROMUSA will also be held during this meeting. Programme for Musa Improvement (PROMUSA)”, article published by the CGIAR

BSV meeting that this is now hap- Within the framework of the Virology pening. working group, a meeting focusing on BSV Since it is not possi- was organized by INIBAP and held in ble to cover all aspects Montpellier from 19-21 January, 1998. The of Musa improvement meeting brought together the most prom- in a short article like inent virologists working on BSV for three PROMUSA, hopefully days of discussions and exchange of this response is not information and ideas. The meeting fo- out of order. Actually, cused on recent advances in the under- a better description standing of the virus and the significance of than response would integrated viral sequences in the Musa be “additional view- genome. Research needs were analysed points”. I know that the and prioritized and current BSV indexing CGIAR frequently in- procedures reassessed in the light of re- vites outside opinions, search results. It was generally agreed that and I thought that the current indexing methods are still the some of my observa- most reliable for detecting a wide range of tions during 28 years BSV isolates, and revisions to the Techni- as a banana and plan- cal Guidelines for the Safe Movement of tain breeder could be Musa germplasm were not considered helpful. necessary. The proceedings of the meeting In examining the will be published shortly by INIBAP. sentiments expressed below, I trust that the Figure 1. FHIA-21 on sale in supermarket in San Pedro Sula, Honduras reader will keep the vital importance of the Genetic Improvement Working Group he above PROMUSA article was pub- subject in mind. Far more than 100,000,000 In relation to the development of molecular lished by the CGIAR in 1997 in the people stand to be directly affected by the markers in Musa as a means of improving TIssues in Agriculture 12 booklet titled decisions concerning allocations of funds for the breeding efficiency, an agreement has “Global Programs: A New Vision in Agricultural development of new disease-resistant banana been signed between CORBANA and Research”. The senior author is the Director of and plantain hybrids. Thus, it is critical that the INIBAP to establish a number of segre- INIBAP. Since May 1994, INIBAP has been right decisions be made. gating populations. These populations will under IPGRI as a member of the CGIAR in- be used in the identification of molecular ternational centers, and INIBAP is responsible for The inadvisability of starting new breeding markers for specific traits and the providing the executive secretariat to PROMUSA. programmes produced would be made available to The stated mission of INIBAP is to increase In my opinion, it would not be advisable to con- research laboratories for molecular marker the productivity and stability of banana and sider starting additional new breeding pro- research. plantain grown on smallholdings for domestic grammes. The reason for this is that the ap- The project, which started in mid-December consumption and for local and export markets. proaches to genetic improvement of Musa are 1997, will run for a period of three years The authors do a very good job of de- quite complicated as compared to those of and four months and will be supervised by scribing the serious disease and pest problems most crops. Dr Jorge Sandoval of CORBANA. on bananas and plantains worldwide, and they There is one key step in the Musa breeding Initial crosses will be made by EMBRAPA, are right when they say that development of process that is of the utmost importance since Brazil to generate one or two segregating disease-resistant hybrids is the most critical all progress in genetic improvement is de- populations which will complement popula- need. I have always wondered why the CGIAR pendent upon it. This key step is the devel- tions available at other institutions. The pop- did not express more interest in banana and opment of agronomically-improved, disease- ulations will be established at CORBANA plantain breeding, and it is encouraging to see resistant diploid parental lines. These improved

INFOMUSA — Vol 7, N°1 PROMUSA I bred diploids are subsequently used as the edible fruit with FHIA-21. The consumer and, assuming good segregation is ob- male parents in cross-pollinations onto seed- benefits by buying FHIA-21, but the farmer ben- tained, will be evaluated for parthenocarpy, fertile triploids for the synthesis of tetraploid efits even more. In all trials comparing FHIA-21 bunch orientation, resistance to black hybrids. (A publication which describes the with False Horn, FHIA-21 has been more than Sigatoka, and agron- advances made in diploid breeding in the FHIA twice as productive than this traditional plantain omic traits. Fusarium wilt evaluation de- programme is: Rowe P.R. and F.E. Rosales. in Honduras. pends on the availability of suitably infested 1996. Bananas and plantains pp. 167-211 in A realistic appreciation of how difficult it has land and will not be part of this project at J. Janick and J. N. Moore (eds.). Fruit Breeding, been to develop improved diploids can be present. Vol. I: and Tropical Fruits. John Wiley & gained by reviewing the published reports of all Sons, New York). other breeding programmes which have been An investment of several million dollars and established in the last 20 years. The most fre- almost 40 years of exhaustive cross-pollinations quently mentioned diploids used in these Nematology Working Group and selections were required to develop the breeding schemes are Calcutta 4 and Pisang The participants in the nematology working genetically-improved diploids in the FHIA pro- Lilin, which are resistant to black Sigatoka, but group have been particularly active in de- gramme. Some of these advanced diploids have very small bunches. These inferior bunch veloping ideas for collaborative projects have already been used as male parents in the sizes are expressed in the tetraploids derived and these will be discussed further at the development of several disease-resistant tetra- from these two diploids. This is in no way in- Cameroon meeting. Ms. Inge Van den ploid hybrids now being cultivated commercially tended as an unkind remark about these pro- Bergh, the INIBAP/VVOB Research Associ- on several thousand hectares. grammes; however, it is intended to show just ate recently appointed to work with VASI in It would be practically impossible to duplicate how extremely difficult, expensive, and time Vietnam has started evaluating Vietnamese these accomplishments in new diploid breeding consuming this diploid improvement scheme germplasm for nematode resistance. Within schemes. All the natural diploids which were has been and is. the framework of this working group, an- collected in Asia and are in germplasm col- The dwarf varieties (which are rightfully other VVOB Research Associate, Thomas lections (and FHIA’s field collection is still the credited with saving millions of lives and for Moens, has recently been appointed to largest in the world) have very inferior features which Dr Norman Borlaug was awarded the work at CORBANA. He will be assisting which include absence of edible pulp, male or Nobel Peace Prize in 1970 for his work in de- with the evaluation for nematode resistance female sterility, unacceptable flavors, and veloping them) and the streak virus- of the segregating populations established agronomically inferior (small) bunches. The un- resistant maize varieties (that greatly increased within the framework of the Genetic im- paralleled accomplishment of my predecessor, the production of this major food crop in Africa) provement working group. Dr D.L. Richardson, was the development of a are two of the outstanding accomplishments of bred diploid, SH-2095, which has outstanding strong, concentrated efforts in international bunch features (very large bunch with long and centers sponsored by the CGIAR. Bananas and parthenocarpic fruits). It took 10 years (1959- plantains are even more important than wheat Fusarium Working Group 69) with the aid of an average of four pro- and corn as domestic food crops in many areas Within the framework of the Fusarium wilt fessional assistants, about 90 laborers, and the of the world. A strong Musa improvement pro- working group, 11 countries have sent evaluations of up to 10,000 diploid plants gramme would be much more effective than se- samples from wilted plants to QDPI, Aus- annually to develop and select SH-2095. veral small programmes for meeting the needs of tralia for isolation and analysis of Fusarium Now, SH-2095 is in the pedigree of prac- banana and plantain farmers and consumers. oxysporum f.sp. cubense (Foc). Vegetative tically all the current improved, disease-resistant compatibility group (VCG) and volatile pro- diploids in the FHIA programme. A good, and breeding vs. biotechnology duction analysis have been completed, while fair, question at this point would be, “Just how In my opinion, the PROMUSA article gives molecular analysis (RAPD-PCR and DAF) valuable have these advanced bred diploids more credit to biotechnology and genetic are still ongoing for those samples where been?”. engineering than has actually been earned. It is VCG’s could not be determined, or which To illustrate this value, let us look at the black not recent developments in biotechnology which had unusual morphology and atypical Sigatoka-resistant FHIA-21 plantain hybrid have allowed some of the barriers to genetic volatile production. The results of the ana- which was developed by using one of these improvement in Musa to be overcome. The de- lysis provide information on the global diver- bred diploids in cross-pollinations onto a seed- velopment of agronomically-improved, disease- sity of Foc and are available from INIBAP fertile plantain. This FHIA-21 plantain hybrid is resistant diploid breeding lines is what has led (Annual report, 1997). One sample from now being grown commercially in Honduras and to surmounting these barriers — just as, more India contained Tropical Race 4 (VCG several other Latin American countries. While than 50 years ago, Dodds in Trinidad correctly 01213/16), the first report of this race of this reference to the cultivation of FHIA-21 is predicted that it would be (Dodds K.S. 1943. Fusarium in India. confined to Latin America, there is now a The genetic system of banana varieties in scarcity of plantains in practically all producing relation to banana breeding. Empir. J. Exp. countries worldwide. This scarcity is because Agric. 11: 89-98). the yield of the universally-cultivated False Horn The Summary of Proceedings and Decisions Global and regional evaluation plantain has been reduced by up to 50% as a of the 27-31 October 1997 International programme result of defoliation by black Sigatoka during the Centers Week in Washington, D.C. has some The majority of the IMTP Phase II evalua- last 20 years. interesting comments about biotechnology. In tion sites have now completed the For this illustration, I went to three super- one place (p. 26), it is said that developments evaluation work and the first results have markets in San Pedro Sula, the second largest in biotechnology “raise important concerns for been received by the programme coordi- city in Honduras, in search of plantains. The the CGIAR, but also present exciting op- nator. The global analysis of the data has first supermarket had only False Horn plantain portunities to accelerate germplasm enhance- already started. Initial results indicate that on sale. These were of poor quality and cost ment via a more strategic research effort”. On the FHIA hybrids are performing well US$0.92 for ten fingers. FHIA-21 was the only another page (p. 7), it is said that, “In Cairo, across a range of environments and dis- plantain available at the second supermarket when we discussed the promise and perils of ease pressures. A first version of the IMTP (Figure 1). The fruit were larger than False Horn biotechnology, we agreed that these questions candidate clone database has been com- (ten fingers weighed 50% more) and the cost must be scrutinized in light of reason, in the pleted and available data is being entered. was US$1.06 for ten fingers. The third presence of evidence, and not governed by The database will eventually contain supermarket had no plantains in stock at all. So, emotion or prejudice”. information on pest/disease resistance/- for a US$0.14 higher price for the same number I am in complete agreement with this latter tolerance and agronomic performance of fingers, the consumer receives 50% more statement by the CGIAR. “In light of reason”, a under specified conditions of improved

II PROMUSA INFOMUSA — Vol 7, N°1 hybrids and other available for review of what led to the historical development There are no known biotech techniques for evaluation by NARS. of almost 100% of the productive and disease- increasing inherent low productivity. However, A number of breeding programmes, in- resistant varieties of practically all food and yield is a very important consideration in the cluding CIRAD, FHIA, IITA and CRBP, fiber crops would reveal that these devel- genetic improvement of plantains. In breeding have agreed to contribute improved hybrids opments were the results of applications of to develop productive, disease-resistant plan- for testing in the next phase of IMTP. Most sound breeding principles by plant breeders. tains by crossing advanced bred diploids onto a of these hybrids have already been re- When biotechnology applications have been seed-fertile plantain clone, black Sigatoka- ceived by the ITC and are presently under- made, these applications have been almost resistant plantain hybrids with double the yield going virus indexing prior to distribution for exclusively to modify plants that had first been of the universally-cultivated False Horn plantain evaluation. improved by plant breeders. have been developed in the FHIA programme. Bananas and plantains have lagged behind The already-discussed FHIA-21 plantain hybrid most other crops in regard to their genetic is one of these developments, but, as good as it improvement for two reasons. The first and pri- is, FHIA-21 has a weakness in that the green New structure for IMTP mary reason is that, until it was proven possible, life of fruit after harvest is relatively short when The International Musa Testing Programme very few people believed that it could be done. compared to that of False Horn. This weakness is now operating within the framework of After all, the varieties which have traditionally requires that harvested fruit be marketed rather PROMUSA. The structure of the programme been cultivated are natural (non-bred) triploid quickly. has therefore been revised to better serve plants with seedless fruit. The other reason is By using a different improved diploid male the germplasm evaluation needs of pro- that, as has been discussed, the development of parent in cross-pollinations onto the same seed- gramme participants. The revised structure improved banana and plantain hybrids was de- fertile plantain that is the female parent of FHIA- also takes into account the needs of the pendent upon first developing the necessary 21, an additional plantain hybrid, FHIA-20, has growing number of national programmes diploid parental lines. In spite of the very capable been selected which also is twice as productive wishing to participate in the programme. work of numerous dedicated scientists, this de- as False Horn. However, FHIA-20 is superior to The main feature of the revised programme velopment of these advanced diploids took FHIA-21 in that the fruit has an excellent long is the introduction of two levels of evalu- several decades because of the inferiority of the green life after harvest comparable to that of ation: natural diploids which were available for begin- False Horn. Like FHIA-21, FHIA-20 is much more • In-depth evaluation: Evaluations that are ning this diploid improvement breeding scheme. resistant to black Sigatoka than False Horn. carried out at the in-depth sites are The “presence of evidence” as cautioned by Bunch characteristics of the FHIA-20 plantain detailed and will include not only disease the CGIAR about biotechnology, will be dis- hybrid as compared to those of the False Horn and pest resistance/tolerance screening cussed further in the succeeding paragraphs. plantain, with no chemicals applied to either for but can also be combined with epidemio- As has been seen with the discussions about control of black Sigatoka, are shown (Figure 2). logical studies on pathogen populations, the FHIA-21 plantain hybrid, the “presence of In addition to its excellent yield, FHIA-20 has studies on host-pathogen relationships for evidence of the value of what is now called exceptional eating qualities, and is considered different races of the pathogen and conventional plant breeding” has already been by many consumers to be superior to False adaptability and yield studies. The evalua- demonstrated once, and it will be confirmed Horn when cooked both green and ripe. This tion protocols used at these sites are again for both plantains and bananas as these bred plantain hybrid is now being grown com- elaborate, requiring time and expertise. discussions continue. mercially in Honduras, and these plantings by The information obtained is primarily In an attempt to show that I am trying to be farmers are expanding rapidly. There is no way useful to breeders and pathologists. objective, I recognize that there have been a few that a new plantain with all the outstanding • Performance evaluations: The majority apparent successes from using biotechnology qualities of FHIA-20 could be developed by of evaluation sites will carry out perfor- applications for modifying certain characteristics using biotech applications. mance evaluations, where the material is of some plants. Some examples are: alterations I know of no biotech “success stories” about assessed for disease resistance, agron- of natural fruit ripening, increasing herbicide re- the introduction of resistance to a fungal di- omic adaptation and stability using a sistance (but this has met with a recent notable sease or a nematode in any crop. Yet, the main simple protocol requiring a minimum of failure in one crop), increasing insect resistance disease and pest problems of bananas, plan- time and expertise. Collaborators at with the Bt toxin (with sporadic success), and in- tains, and cooking bananas are, in order of these sites can select the clones they creasing virus resistance by introduction of virus current relative importance: black Sigatoka (a wish to evaluate based on information coat protein into the genome of certain plants. fungus), the burrowing nematode, and Panama provided by INIBAP, according to local Thus, I concede that biotech may have some ap- disease (a fungus). In addition, the resistances needs and conditions. The information plications for some traits in some plants. But now, to black Sigatoka, Panama disease, and the obtained is primarily useful to extension let us return to the discussion of biotech applica- burrowing nematode which have been identified agents and farmers. tions for improvement of bananas and plantains. in natural banana clones are controlled by Revised protocols for the two levels of evaluation have been developed by the Fusarium and Sigatoka working groups and are being published by INIBAP. Both types of evaluation sites provide feedback information on the agronomy, pathology and adaptation of improved varieties to INIBAP. This in- formation is compiled in a database, which will be linked the Musa Germplasm Improve- ment System (MGIS) database. Information is then fed back to the breeding programmes and is also available for other NARS to assist in their choice of appropriate, well charac- terised varieties for testing. If you would like to receive more information about the IMTP, or are interested in participating in the programme, please contact the IMTP Coordinator at INIBAP Figure 2. Typical bunch characteristics of the universally cultivated False Horn plantain (center) as compared to those of the black Headquarters, Montpellier France. Sigatoka-resistant FHIA-20 plantain hybrid (on both sides) in Honduras when no chemicals are applied for control of black Sigatoka

INFOMUSA — Vol 7, N°1 PROMUSA III several genes with an additive effect. Thus, diploid is a preferred parental line in these since it would be practically impossible to studies because of its resistance to race 4 of identify and isolate so many genes, biotech is Panama disease and to the burrowing nem- doubly limited in its applications to disease and atode. However, PJB is so highly sterile that it pest resistance in bananas and plantains. (An cannot be used in cross-pollinations for this excellent popular-style article about biotechnol- purpose. ogy regarding its applications to crop improve- My suggestion would be that the funds avail- ment is: Simmonds N.W. 1997. Pie in the sky. able for “molecular marker studies” could, The Planter 73(860): 615-623, November 1997. instead, be put to a very valuable practical use I would gladly send a copy of this article to by greatly increasing the numbers of bunches interested readers upon request.) pollinated in breeding for the development of The fact remains that practically all cultivated dwarf, black Sigatoka-resistant plantain hybrids. food and fiber crops have been genetically (The FHIA-20 and FHIA-21 plantain hybrids improved, and these improvements have been described earlier are tall plants like False Horn, made by plant breeders. As shown by FHIA-20 and thus, as is False Horn, susceptible to blow- and FHIA-21, which were described above, and downs by strong winds. A plantain hybrid with a the FHIA-23 banana hybrid, bananas and plan- shorter plant height would significantly reduce tains have now joined that list. FHIA-23 is a these losses caused by winds, which, in turn, tetraploid dessert banana which was derived would lessen the periodic lengthy interruptions from crossing the SH-3362 bred diploid onto in availability of fruit.) the Highgate dwarf mutant of Gros Michel (Gros Michel was the original and only export Conclusions banana before it was destroyed by race 1 of I know of no more urgent food security problem Panama disease). This new banana hybrid is in the world than that confronting the plantain resistant to race 1 of Panama disease and and farmers and consumers in tolerant to black Sigatoka. More than anything Africa. And, I know of no plant disease that else, these FHIA-20, FHIA-21, and FHIA-23 currently adversely affects more people (about commercial hybrids are indicator plants. They Figure 3. Bunch characteristics of two diploids illustrate one of 70,000,000 in West and Central Africa and at show that disease-resistant, productive hybrids the major accomplishments in the FHIA breeding programme. least 30,000,000 in Latin America) than black Pisang Jari Buaya (left) is a natural diploid clone from Asia and of bananas and plantains which are preferred SH-3362 (right) is a bred diploid, a direct descendant of Pisang Sigatoka on plantains. When black Sigatoka by farmers and consumers can be bred, and Jaru Buaya by way of cross-pollinations and selections spreads throughout East Africa, as is already are indicative that even better hybrids can now happening, the production of the main food (the be expected to be forthcoming — if these con- development of SH-3362 are not possible black Sigatoka-susceptible highland cooking tinued efforts in this successful programme are by gene manipulation with biotechnology bananas unique to this area of Africa) for adequately supported. techniques. 20,000,000 people is projected to be reduced As emphasized earlier, and it is of such by 50% of that currently being produced. Race paramount importance that it merits this addi- The questionable value of molecular 4 of Panama disease is already a serious tional emphasis, all progress in Musa breeding markers problem in some countries, and if it arrives to has been dependent upon the development of An action plan advocated in PROMUSA is the Latin America, it would destroy the export ba- advanced diploid breeding lines, which serve identification of molecular markers. In my nana. The losses of jobs, foreign exchange, and as the male parents in the pollinations of opinion, this activity should be given a low an important tax base, which would go along naturally occurring triploids (the cultivated priority. Based on my experience, a knowledge with such a destruction of the export banana, bananas and plantains are examples of tri- of “molecular markers” would not have been would devastate the economies of several ploids, but not the ones used in breeding useful in the development of any of the ad- countries. because they do not produce seeds). With one vanced breeding lines in the FHIA programme, However, there is hope because these needs exception, diploids are not cultivated for nor would such markers have aided in the de- can be addressed by breeding disease-resistant commercial production, and their main value is velopment and selection of the several pro- hybrids. In fact, breeding is the only practical as sources of disease and pest resistances in ductive, disease-resistant banana, plantain, and solution, and substantial progress has already breeding programmes. The diploid with the cooking banana hybrids now being grown been made in this regard (some examples of best bunch characteristics of all the many commercially. this progress were pointed out in the above diploids which have been collected in germ- In my career as a plant breeder, I have had descriptions of the development of FHIA-20, plasm collection expeditions is Pisang Jari the good fortune of knowing about 25 very FHIA-21, and FHIA-23). Buaya (PJB). PJB is almost sterile (and has no successful breeders of different crops and being The authors of PROMUSA are to be com- pollen) for breeding purposes, but has been exposed to their approaches to genetic im- mended for bringing the great needs for the found to be resistant to race 4 of Panama di- provement. I do not know of an example in which development of disease-resistant bananas and sease and the burrowing nematode, and molecular markers would have been useful to plantains to the attention of the CGIAR. In this tolerant to black Sigatoka. One of the most these plant breeders in their development of response, I have attempted to emphasize the significant accomplishments in the FHIA pro- varieties which became the standards of excel- advantages of supporting a proven breeding gramme has been the development of the lence for their chosen crops. programme to accomplish these objectives. I SH-3362 bred diploid which has PJB in its There are unavoidable barriers to estab- trust that the arguments and illustrations pre- pedigree (Figure 3). In addition to its out- lishing segregating populations for subsequent sented will be useful towards determining the standing agronomic (large bunch) qualities, identification of molecular markers in Musa. For best courses of action in this very important SH-3362 is also resistant to races 1 and 4 of any reliable genetic study, homozygous par- Musa improvement project. Panama disease, and is tolerant to the ental lines are a must, and such parental lines Phil Rowe burrowing nematode and black Sigatoka. are practically non-existent in Musa. Plus, La Lima, Honduras SH-3362 has been an extremely valuable populations from backcrosses to each parent March 17, 1998 parent in further cross-pollinations, both in the and an F2 population would have to be pro- development of breeding lines with multiple duced and evaluated, and it is practically Phil Rowe is the Head of the Banana and disease resistance and in the synthesis of the impossible to do this because of sterilities (both Plantain Programme of the Fundación Hondureña FHIA-23 tetraploid, which was described in the male and female) in the most indicated parental de Investigación Agrícola FHIA), Apartado postal preceding paragraph. Accomplishments like the lines for these studies. For example, the PJB 2067, San Pedro Sula, Honduras.

IV PROMUSA INFOMUSA — Vol 7, N°1