The Distribution of Benefits from Public International Germplasm Banks: the Case of Beans

The Distribution of Benefits from Public International Germplasm Banks: The Case of Beans in Latin America

Oswaldo Voysest,1 Nancy Johnson2, Douglas Pachico3

January 14, 2002

Abstract

The unrestricted international flow of genetic resources from international gene bank collections is perhaps one of the greatest impacts of international agricultural research. This paper examines the distribution of benefits from bean genetic resources across countries in Latin America. The genealogy of commercial bean cultivars released since 1976 is analyzed to calculate for each country the source of the genetic resources used commercially in that country. All countries are shown to be heavily dependent on imported genetic resources for their commercial cultivars. From information on the economic impact of improved bean varieties, the share of economic productivity benefits associated with imported germplasm by country of origin is calculated. The benefits received by each country from improved bean germplasm are compared with the contribution of that country’s germplasm to other countries. Some of the patterns in the flow and use of genetic resources are analyzed.

1. Introduction

Large increases in crop productivity have been achieved through the sciences of plant genetics and breeding, including breakthroughs like hybrid maize and semi dwarf wheat and rice. Improved crop varieties associated with the International Agricultural Research Centers (IARCs) have generated economic benefits worth billions of dollars (Evenson and Gollin, 2001). These benefits derive from the application of advanced scientific methods in plant breeding, however they also depend on the availability of raw materials, genetic resources with desirable traits that can be combined to produce a plant more productive than its parents. It has been estimated that up to one half of the gains in U.S.A. agricultural yields from 1930-1980 can be attributed to genetic resources (Office of Technology Assessment 1987).

Much of the germplasm used for crop improvement comes from germplasm collections that are the result of decades of collection and conservation of materials from all over the world. In the past, collection and exchange of germplasm was relatively unregulated except for basic phytosanity requirements. Recently, interest in intellectual property rights to genetic resources has led to international conventions that assign ownership of materials to their country of origin and regulate their exchange. While it is important that

1 Agronomist, formerly with CIAT Bean Program (o.voysest.cgiar.org) 2 Economist, Impact Assessment Project, CIAT ([email protected]) 3 Economist and Director of Strategic Planning, CIAT ([email protected]) countries recognize the economic value of their genetic resources, there is a danger that unilateral attempts to appropriate these benefits could end up restricting the flow of germplasm. The consequences of an interruption in the flow of genetic materials would depend on how important “imported” materials are to the production of improved varieties released by individual countries.

This paper documents and evaluates the importance of inter-country gene flows to Latin America and the Caribbean by analyzing the genealogies of the improved bean varieties produced between 1975 and 2000 using materials from CIAT’s bean germplasm collection. The national origins of the genebank accessions that appear in the pedigrees of improved varieties are identified, and the patterns of germplasm exchanges analyzed. The economic value of gene flows is calculated by apportioning the estimated benefits of the improved varieties—in the form of the value of increased production—among their ancestors. The results of the analysis show how individual countries both benefit from and contribute to the flow of germplasm from the CIAT gene bank.

The paper is organized as follows. Section 2 describes the germplasm collection, and the exchange and improvement efforts that led to the release of improved varieties. Section 3 presents the results of a genealogical analysis of the varieties improved varieties. In section 4, the economic benefits associated with the exchange of genetic material are calculated using available information on the adoption and impact of improved varieties. Section 5 discusses the main findings and their implications for research and for genetic resource conservation and use.

2. Conservation and use of bean genetic resources in Latin America

Common bean is a noncentric crop, with multiple domestication sites throughout Middle and Andean South America. The genetic resources of the common bean consist of two major gene pools, one Mesoamerican and one Andean. Wild ancestors and common bean land races from these two sources are the founding genetic base of today’s cultivated beans, which consist of six main races based on geographic origin and seed size.4

Genetic heterogeneity in cultivated beans exists at two levels (Adams, 1977). In Latin America, the first and upper level of the bean mosaic diversity would be the polymorphic expression of beans for plant and seed characteristics. The other level of diversity is formed by the diversity among cultivars within each production region. Patterns of variation at the first level have been thoroughly analyzed (Gepts et al, 1986); Singh, 1989, 1992, Singh et al, 1991; Voysest and Dessert, 1991).

A significant portion of bean genetic diversity is in the custody of the Genetic Resources Unit (GRU) of the International Center for Tropical Agriculture (CIAT) where, within certain limitations, its flow and use can be monitored. Another large portion of the

4 Chile, Nueva Granada and Peru are large seeded Andean races. Durango and Jalisco are medium seeded Middle American races, and Mesoamerica is a small-seeded Middle American race.

2 common bean genetic resources has not been yet properly catalogued. This paper deals with the designated germplasm, i.e. the bean genetic resources which CIAT holds in-trust for the world’s farming community and that are kept in the GRU with a coded G identification number.

The CIAT-GRU, or germplasm collection, currently has 31,880 accessions of Phaseolus beans designated to FAO. It is the largest collection of bean genetic resources in the world, accounting for 15 percent of total accession in ex situ conservation. In addition to collected material including Phaseolus species, wild populations of Phaseolus vulgaris, unimproved landraces and obsolete varieties, the collection also includes pure lines, modern elite varieties, and breeding lines. The collection includes materials from 65 countries. The national origin of an accession is defined as the country where the material was collected or developed. Main contributors to the gene bank include Mexico (4706 accessions), Peru (2,361 accessions) and Guatemala (2,209) and Colombia (1,380). The USA has contributed over 10,000 accessions, though in many cases the USA is the source rather than the country of origin.

Since 1973, over 307,493 accessions have been sent out by the germplasm bank to 96 countries to be used in bean research and development programs. The top five recipients were Colombia (16.2%), USA (9.1%). Guatemala (8.6), Brazil (6.2%) and Mexico (6.0%) (CIAT-CRU, 2001). Excluding materials used in CIAT, 60% of distributed materials went to national research systems (NARS) and 33% went to universities, with the remainder going to NGOs, commercial companies, individual farmers, other gene banks, and other CG centers (Debouck, 2001). Forty four percent of the materials distributed by the germplasm collection were used for breeding. The remainder were used for basic and applied research (41%), agronomy (14%) and training purposes (1%). CIAT scientists are much more likely to use materials for breeding than are scientists from external institutions.

3. Contribution of the germplasm bank to varietal development in Latin America

Released varieties and their pedigrees were obtained from published CIAT sources (Voysest, 1991, 2000), circulars and bulletins, the CIAT Bean Project database (Rodriguez et al.), personal communications with bean breeders and other sources (Mc. Clean and Myers, 1990). Pedigrees of the cultivars were traced back to ancestors in the gene bank with G identification numbers or genotypes for which no further genealogical relationships could be discovered. For calculating the genetic contribution of each hybrid genotype we assumed that a cultivar received half of its genes from each parent and that genetic drift and selection did not cause significant deviations from the expected relationships. It was assumed that parents used in crosses were homozygous and homogeneous.

From the accessions of Phaseolus beans designated to FAO and kept in custody in the CIAT –GRU, 193 were used to develop 243 cultivars which have been released in 18 countries of Latin America between 1976 and 2000 (Table 1). The total number of varieties released in LAC during the period was 411. Of these, 243 are known to have made use of the CIAT germplasm collection while the remainder were developed directly

3 by national programs without input from CIAT or the gene bank. This study focuses on the 243 CIAT genebank-related varieties and the role of the genebank in their development.

Twenty-eight genebank accessions were released directly as cultivars in Latin America. Because some were released in more than one country, they resulted in 38 varieties (Table 1). Of these 38 varieties selected directly from genebank accessions, 14 belong to the Andean gene pool and 24 to the Mesoamerican gene pool. Designated accessions from Brazil, Colombia, Mexico and the USA were the most used for selecting new cultivars. Peru and Bolivia were the main users of designated accessions to select these new cultivars.

Although the genebank was an important direct source for selecting new cultivars, in Latin America its greatest contribution lies in its use by breeders to develop 205 new varieties through hybridization. A total of 179 genebank accessions were used as parents in these crosses (Table 2). Argentina has released the largest number of crosses using material from the genebank; 28 varieties were released that included 66 genebank parents from 20 countries. Brazil released 34 varieties containing 55 genebank accessions from 17 countries. Countries obtained genebank parents in two ways. In some cases national breeders directly requested materials from the genebank and used them as parents in their own breeding programs. In most cases, genebank materials were obtained indirectly through the receipt of bred material from CIAT breeders who had used genebank parents. In this second group of cases, genebank parents originating in a given country were sometimes used in crosses that were sent to that same country. In this way, genebank parents were sometimes sent back to their countries of origin. In this paper, the use of genebank parents in their country of origin is not treated as a contribution of the genebank because in principle, these material could have been locally available to national breeders. Thus, this paper somewhat underestimates that actual contribution of the genebank.

The total number of countries whose germplasm has been included in released varieties is one way of assessing a given country’s use of foreign designated germplasm. Another is to look at what percent of the total genetic content comes from imported genebank accessions. Only in Colombia and the Dominican Republic did local germplasm contribute half or more of the genes of the new varieties (Table 3). Again, much of this “local” contribution actually came from CIAT bred material that used genebank parents originating the respective countries, so most of the “local” content is in fact actually associated with the genebank. At the opposite extreme are Venezuela, Bolivia, Panama, Cuba and Haiti, where locally-originating genebank parents account for less than 10 percent of genetic content of released varieties.

In countries where large and medium seed sizes (Andean gene pool characteristics) are preferred, such as Colombia, Ecuador, Peru, Chile and the Dominican Republic, germplasm from Colombia was the most important source of genes (Table 3). In regions where small seeds are preferred, germplasm from Mesoamerica made the greatest genetic contribution. The presence of USA germplasm and the absence of Peruvian germplasm as

4 important sources of genes was quite a surprise. Peru is a major center of bean genetic diversity, however this diversity appears not to have been widely used in the production of improved varieties. In contrast, germplasm from the USA makes up a small share of diversity in the bean genebank, but it has been widely used in developing new cultivars.

Among the 31 countries that contributed the 179 genebank accessions used to develop the 205 varieties analyzed in this study, 11 supplied close to 80% of the genes. The contribution of the remaining 19 countries was relatively small. Colombia was the country with the greatest genetic contribution, contributing a total of 17 percent of the material in released varieties in other LAC countries (Table 3). This was particularly evident in the countries where Andean–type varieties are preferred. Mexico, Costa Rica and El Salvador were other important donors particularly in the countries where Mesoamerican type beans are grown. The contribution of germplasm from the USA was evident in all countries of Latin America, particularly important in the countries where large seeded varieties are grown.

4. The economic value of germplasm exchange

The previous section documented the importance of germplasm exchange among countries in the development of improved varieties in Latin America. No country can claim complete reliance on its own genetic resources, and all countries made significant use of the genetic resources of other countries. The genetic contribution of the pool of 31 donors via 193 germplasm accessions is impressive. Even Colombia, Mexico and Brazil, countries with strong breeding programs and abundant genetic resources, show a significant use of foreign germplasm.

In order to generate economic value, it is not sufficient that genetic materials be incorporated into released varieties. Those varieties must also be adopted by farmers and contribute to increases in productivity. Available data on the adoption and impact of CIAT genebank varieties show that they have achieved relatively high use in the LAC region (Johnson et al, 2001; Table 4). An estimated 49 percent of bean area in LAC was planted to CIAT genebank varieties in 1998, ranging from 16 percent in Peru to 85 percent in Costa Rica. The yield gains associated with these varieties average 210 kilos per hectare, against an average per hectare yield of between 500 and 1000 kilos.5 In 1998, the annual value of increased production associated with the varieties was estimated to be US$177.5 million, and the cumulative value of increased production due to improved varieties between 1976 and 1998 was over a billion dollars.

We do not have complete data on impact by variety, however through expert opinion we were able to eliminate the released varieties that were known to have experienced little acceptance by farmers. The analysis presented in previous sections was re-done for the remaining economically-important varieties. If we assume that all genes contribute equally to the economic value of a variety, then on a regional basis, 72 percent of the

5 Reported yield gains are net gains associated with improved varieties. Yield gains associated with increased input or other changes in management practices are not included.

5 benefits of improved varieties—US$827 million—can be attributed to “foreign” germplasm (Table 6)6. The reason it is so high that many of the largest bean producing countries such as Brazil and Argentina are big users imported germplasm from the CIAT genebank. To estimate the value of bilateral exchanges of germplasm, we can allocate a country’s benefits from improved varieties among the countries whose germplasm it used (Table 5). For example, the proportion of Brazil’s benefits from improved varieties that can be attributed to Costa Rica would be equal to the proportion of Costa Rican germplasm in the varieties released and adopted in Brazil.

According to the results, Mexico has made the biggest economic contribution to varietal development in other countries (USD168 million), followed by El Salvador (USD121 Million), Costa Rica (USD103 million) and the USA (USD 99 million) (Table 6). European and African countries also made important contributions to germplasm development in LAC. Among the countries for which data were available, Brazil was the biggest beneficiary of international germplasm exchange (US$538 million) followed by Argentina (US$115 million) and Guatemala (US$53 million). Six countries are net beneficiaries and six are net donors. Brazil and Argentina continue to be the largest net beneficiaries of international germplasm flows. El Salvador, Colombia, and Costa Rica are the biggest net contributors.

One of the principle motivations for assigning rights to genetic materials is to reward those who developed and conserved them. The hypothesis is that collection of royalties from the use of germplasm could be a source of income for many developing countries. The results of an empirical analysis done at the regional level show that while this may in fact be true, many of the transfers would occur between developing countries rather than from developed to developing countries (Pachico, 2001). In most cases, the production benefits that countries receive from using imported germplasm outweigh any potential benefits they could obtain via collection of royalty. We can do a similar analysis here using data from the country level. Assuming a 10 percent royalty payment to countries on the basis of the value that their genetic materials generate in other countries, only Colombia would earn more through royalties than through the productivity benefits associated with their own use of imported germplasm. As will be discussed below, this result is likely misleading because much of the germplasm attributed to Colombia is actually associated the international genetic improvement programs that have operated in Colombia for decades.

5. Conclusions and discussion

CIAT’s bean genebank is used for a wide variety of purposes both inside and outside CIAT. One of the principle uses is as a source of genetic materials for breeding. Between 1976 and 2000, of the 411 bean varieties released in Latin America, 243 contained materials from the CIAT genebank. The 193 genebank accessions used to develop these varieties came from 31 countries from LAC, North America, Europe, and

6 In this analysis, all credit for improved productivity is implicitly assigned to the genetic resources. In reality, the input of the conservation and breeding programs must also be acknowledged. This issue will be addressed in future work.

6 Africa. The use of foreign genetic material is widespread in LAC, demonstrating the importance of international exchange of germplasm. Colombia and the Dominican Republic were the only countries whose released vareieties contained more than half of their genes from local sources. Eleven of the 18 countries studied got more than 70 percent of the genes in released varieties from other countries. Colombia was the biggest contributor of germplasm to other countries, followed by Mexico, Costa Rica, and El Salvador.

These international gene flows are associated with significant economic value in the form in increases in bean production due to improved varieties. Over 70 percent of the total value of increased production from use of improved beans in LAC can be attributed imported germplasm. Brazil and Argentina are the biggest beneficiaries of economic benefits from imported germplasm, not only because their use of imported genetic material is high but also because they are large bean producers. Germplasm from Mexico, El Salvador, Costa Rica and the United States generated the highest economic benefits in other countries. While these contributions to other countries are important, in general countries benefit more from the productivity benefits of their own use of foreign germplams than they would from charging royalties on use of their germplasm in other countries. Colombia is the only exception.

There is predominance in the use of gerrmplasm belonging to the Mesoamerican gene pool as opposed to the Andean. Even though this is consistent with the fact that a large number of hectares are planted to the commercially-important Mesoamerican varieties, we expected to see a more significant contribution of the Andean germplasm. Especially surprising is the absence of Chilean and Peruvian germplasm. Andean germplasm is a major source of diversity and constitutes a large portion of the germplasm collection. Similarly unexpected was the large contribution of germplasm from the USA. Again, the conservative breeding strategies that most breeders have followed might explain this. Excessive reliance by breeders on well-known sources for disease resistance, for example, might be one of the causes. The USA germplasm used for example is a well- known source of resistance to common bacterial blight and anthracnose a serious problem in bean fields of most growing regions.

The apparent low use of Peruvian germplasm is likely also related to the way genetic resources are defined in this study. This analysis focused on the accessions as they are catalogued in the CIAT germplasm bank. However among the 179 genebank accessions used to develop improved varieties are 32 accessions that are themselves known to be of hybrid origin developed prior to the estabishment of the germplasm collection. Thirteen of these 32 are considered Colombian varieties, 9 are from the USA and the rest from 5 other countries. All of the US varieties derived from germplasm that was also classified as being of US origin, however the hybrid varieties that are attributed to Colombia in the genebank actually include genetic material from Peru, Italy, Japan and the USA. The fact that these accessions are attributed to Colombia results in an overstatement of the importance of Colombia’s genetic resources—though the scientific contribution made in Colombia to develop these varieties deserves recognition—and an understatement of value of resources from other countries such as Peru. The genealogies of these hybrid

7 varieties will be examined in future work. While this will help refine the results presented in this study, it will not affect the finding regarding the fundamental importance of international exchange of germplasm in improving bean productivity.

The issue of national origin of genetic resources raises a more complex question about how to apportion rights and responsibilities regarding collection, conservation and exchange of materials that have evolved over time and space. For example, this study found Colombia, El Salvador, Costa Rica to be important contributors to genetic diversity in other countries, yet there is no scientific evidence to support the domestication of beans in those countries (Gepts P, Osborn TC, Rashka K, and Bliss FA. 1986; Gepts P, and Debouck DG. 1991; Khairallah MM, Sears BB, and Adams MW. 1992; Debouck DG; Beebe S, Rengifo J, Gaitan E, Duque MC, and Tohme J. 2001). Conversely, recent evidence suggests that Bolivia may have been an important domestication site (Beebe et al, 2001), yet no Bolivian accessions were used in the improved varieties studied here. The implication is that since important modifications and/or uses of cultivars often occur outside their centers of origin, management of genetic resources is of global concern.

Another unexpected finding of the study is the relatively low use of genetic diversity in the development of new varieties. Only 179 of the over 30,000 accessions in the germplasm collection were ultimately used by breeders in the development of new varieties. In the case of rice, a crop with a much longer history of breeding and improvement, 833 landraces from a collection of over 80,000 have been used in the production of 1709 released varieties (Evenson and Gollin, 1997). As expected, use of genetic diversity in improved beans appears to be lower than in rice, however in both cases there appear to be significant amounts of unexploited genetic diversity available.

The entire 31,880 materials in the genebank have in fact been used. Essentially all of the materials in the genebank have been evaluated at one time or other for a wide variety of characteristics including resistance to a wide array of diseases and insects, capacity for biological nitrogen fixation, adaptation to poor soils, drought tolerance, and nutrient content among others. Most genebank materials have undergone multiple evaluations for different traits. The small number of genebank materials that have been used as parents in breeding come from the most outstanding perfomers in these extensive evaluations. In the absence of a large genebank, there would be no way of systematically identifying the precious few cultivars with the outstanding characteristics of exceptional parents, nor would there be any guarantee that these outstanding parents would have been conserved.

One reason for the relatively limited use of diversity in beans is that breeders have focused their efforts on breeding for certain commercially-important grain types, while the collection contains many accession from other types. Moreover, conventional breeding techniques have limited the ability to exploit the full range of available genetic diversity. Conventional sexual crosses among different pools of bean genetic diversity often do not yield viable progeny. This has severely constrained breeders’ ability to bring known desirable traits into popular commercial types. New methods of more targeted gene transfer through genetic engineering could help to expand a broader use of genetic resources. Likewise, conventional field based phenotypic evaluations do not always give

8 a precise indication of the genetic potential of a material. New biotechnology-based approaches to gene discovery should facilitate the identification of useful genes that previously could not be directly observed. This too may facilitate a broader use of the existing wide genetic diversity to improve crop productivity.

9 References

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10 Office of Technology Assessment (1987) Technologies to Maintain Biological Diversity. U.S.A. Government Printing Office, Washington, D.C.

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11 TABLE 1. Number of varieties released in Latin America during the period 1976-2000 Country/Region All Varieties CIAT Genebank Varieties All Selections Crosses Brazil 110 37 3 34 Mexico >50 11 - 11 Argentina 31 29 1 28 Chile 17 6 1 5 Guatemala 17 14 1 13 El Salvador 8 6 - 6 Honduras 10 10 1 9 Nicaragua 15 14 - 14 Costa Rica 12 12 - 12 Panama 10 8 3 5 Cuba 18 17 3 14 Dominican R. 11 4 1 3 Haiti 4 4 - 4 Bolivia 17 16 7 9 Colombia 18 14 1 13 Ecuador 15 10 2 8 Peru 42 25 13 12 Venezuela 6 6 1 5 Total 411 243 38 205

TABLE 2. Origin and number of genebank accessions in the development of 205 CIAT cultivars through hybridization in Latin America. 1976-2000 No. and origin of the genebank accessions Released cultivars Country No. of accessions No. of donor countries Argentina. 28 66 20 Bolivia 9 50 15 Brazil 34 55 17 Chile 5 11 8 Colombia 13 25 6 Costa Rica 12 38 12 Cuba 14 34 11 Dominican R. 3 11 6 Ecuador 8 14 6 El Salvador 6 26 13 Guatemala 13 29 11 Haiti 4 17 10 Honduras 9 23 11 Mexico 11 48 13 Nicaragua 14 24 10 Panama 5 21 11 Peru 12 28 13 Venezuela 5 18 8

12 TABLE 3. Total genetic contribution of foreign germplasm, and contributions of ten most important donor countries to the development of improved varieties Proportion Country/ COL MEX CRI SLV USA GUA BRA HON NIC DOM Region of foreign genes Brazil 0.764 .085 .109 .136 .135 .101 .061 NA 0 .043 .007 México 0.568 .100 .083 .057 .049 .043 .246 .026 .026 .033 Central America Costa Rica 0.841 .089 .293 .282 .035 .083 .021 .022 .035 .024 El Salvador 0.815 .042 .137 .152 .031 .016 .001 .138 .062 .083 Guatemala 0.749 .091 .209 .166 .101 .024 .067 .060 .020 .007 Honduras 0.726 0 .212 .111 .076 .036 .010 0 .115 .087 Nicaragua 0.766 .009 .145 .150 .219 .013 .056 0 .152 .018 Panama 0.95 .375 .176 .023 .023 .065 .004 .063 .027 .035 .020 Andean Zone Bolivia 0.973 .151 .138 .058 .005 .166 .078 .205 .009 .037 .060 Colombia 0.255 .048 0 .005 .106 0 .058 0 0 0 Ecuador 0.625 .438 .031 0 0 .062 0 .063 0 0 .031 Peru 0.631 .199 .060 .026 0 .104 .005 .057 0 .007 .042 Venezuela 1 .125 .050 .150 .200 .100 .250 0 0 .025 0 Southern Cone Argentina 0.905 .159 .96 .073 .084 .172 .068 .091 .014 .010 .011 Chile 0.7 .200 .025 .075 .025 .100 .025 0 0 0 0 The Caribbean Cuba 0.929 .072 .205 .131 .125 .034 .064 .049 .092 .110 .025 Dom. Rep. 0.5 .229 .021 .125 .021 .021 .083 0 0 0 Haití 0.938 .125 .113 .164 .180 .010 .129 0 .137 .035 .035

13 Table 4 Impact of CIAT –related bean varieties in selected countries of Latin America, 1998 Area Percent in Yield gain Incremental Value of planted to CIAT- of over local production increased beans genebank varieties (Tons) production (Has) varieties (t/h) (US$) COSTA RICA 39,000 85 0.10 3,290 1,447,432 GUATEMALA 122,780 40 0.23 11,272 7,434,601 NICARAGUA 153,720 30 0.23 10,399 3,529,816 HONDURAS 83,000 45 0.11 4,152 1,409,295 EL 85,000 25 0.29 6,059 2,325,588 SALVADOR PANAMA 11,000 40 0.25 1,105 727,021 BRAZIL 3,307,760 51 0.17 279,211 125,873,960 ARGENTINA 285,000 77 0.24 53,270 28,818,153 COLOMBIA 138,022 10 0.20 2,787 1,198,827 BOLIVIA 11,640 82 0.25 2,375 920,799 ECUADOR 61,520 20 0.13 1,683 889,363 PERU 73,334 16 0.35 4,001 2,878,131 LAC Total 4,371,776 49 0.21 379,604 177,452,986 Source: Johnson et al, 2001

Table 5 Value of economically-important genebank accessions to selected countries in Latin America, by source (1990 US$million) Recipients Region BR MX CR ES GU HO NI CO EC PE VE AR DR US Brazil NA 108.87 69.77 82.04 38.34 0.00 42.17 50.60 12.27 7.67 13.03 0.00 7.67 64.40 Mexico 0.00 NA 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Central America Costa Rica 1.10 7.84 NA 5.77 4.42 1.21 1.85 0.29 0.00 0.00 2.78 0.00 1.28 1.89 El Salvador 0.02 2.66 2.95 NA 0.31 2.68 1.20 0.82 0.00 0.00 1.01 0.00 1.61 0.60 Guatemala 6.96 19.54 8.62 9.42 NA 2.69 2.14 1.50 0.00 0.00 0.16 0.00 0.24 2.22 Honduras 0.00 4.55 2.38 1.63 0.21 NA 2.47 0.00 0.00 0.00 0.34 0.00 1.87 0.77 Nicaragua 0.00 6.58 6.76 6.62 0.18 8.17 NA 0.64 0.00 0.00 0.32 0.00 1.27 0.95 Panama 0.50 1.38 0.18 0.18 0.03 0.21 0.28 2.95 0.00 0.00 1.00 0.00 0.16 0.51 Andean Zone Bolivia 1.08 0.45 0.18 0.00 0.18 0.00 0.13 0.54 0.00 0.00 0.00 0.00 0.00 0.31 Colombia 0.81 0.80 0.00 0.00 0.00 0.00 0.00 NA 0.00 0.40 0.00 0.00 0.00 1.41 Ecuador 0.83 0.42 0.00 0.00 0.00 0.00 0.00 3.32 NA 0.00 0.00 0.00 0.42 0.00 Peru 0.00 1.21 0.31 0.00 0.00 0.00 0.00 3.37 1.21 NA 0.31 0.00 0.31 2.45 Venezuela 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 NA 0.00 0.00 0.00 Southern Cone Argentina 11.17 11.57 12.11 15.21 6.73 3.10 0.67 19.11 2.29 0.00 4.58 NA 0.27 23.28 Chile 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Caribbean Cuba 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Dominican R. 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 NA 0.00 Haiti 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00

14 CAR=Puerto Rico and the Antilles (ATL)

Table 6 Value and distribution of cumulative benefits from improved bean varieties associated with CIAT genebank, 1970-1998 (in US$1990 millions) Beneficiary Cumulative Value Value Net benefit Country value of contributed to received from CIAT benefits LAC via from other gene bank 1970-1988 genebank countries via (not including genebank own country) Brazil 766.72 22.47 538.24 515.76 Not 167.87 Na Mexico available Central America Costa Rica 35.63 103.27 28.75 -74.52 El Salvador 19.41 120.87 15.78 -105.09 Guatemala 79.12 50.40 53.49 3.09 Honduras 21.44 18.05 15.57 -2.49 Nicaragua 45.37 50.91 31.49 -19.42 Panama 7.86 0 7.37 7.37 Andean Zone Bolivia 2.87 0 2.87 2.87 Colombia 12.93 83.13 3.43 -79.70 Ecuador 8.31 15.77 4.99 -10.78 Peru 19.57 8.07 10.39 2.32 Not na Venezuela available 23.53 Southern Cone Argentina 134.59 0 114.94 114.94 Other Countries/ Regions Dominican Rep. 15.09 USA 98.80 Canada 2.29 Europe 27.55 Africa 12.81