During six years exploration of plant genetic resources in Cuba, 328 bean accessions have been collected, ..... On the other hand, historical data show that.
Euphytica 57: 1-8, 1991. © 1991 Kluwer Academic Publishers. Printed in the Netherlands.
Origin, diversity and utilization of the Cuban germplasm of common bean (Phaseolus vulgaris L.) L. Castifieiras ~, M. Esquivel 1, L. Lioi 2 & K. Hammer 3 t lnstituto de Investigaciones Fundamentales en Agricultura Tropical (INIFAT), Santiago de las Vegas, Cuba; 2Istituto del Germoplasma, 70126 Bari, Italy; ~lnstitutfiir Genetik und Kulturpflanzenforschung (IGK), D-0-4325 Gatersleben, Germany Received 11 April 1991;accepted5 September 1991
Key words: Phaseolus vulgaris, dry bean, genetic resources, variation, multivariate analysis, Cuba Summary During six years exploration of plant genetic resources in Cuba, 328 bean accessions have been collected, demonstrating the rich germplasm of this crop. A group of 96 representative accessions was studied for 34 morpho-agronomical characters. Principal Component Analysis and Correspondence Factor Analysis were used to compare accessions. Basic seed colours are black (53%), red (25%) and white (0.3%). Probably first bean introductions were large-seeded cultivars with phaseolin type T, which could have arrived with Taino Indians from South America. Races with small black seeds and phaseolin type S are prevailing in Cuban bean germplasm and could have arrived from Mexico, via the Northern coast of South America and the Antilles arch and reintroduced directly from Mexico after the Conquest. The economic importance of Cuban bean germplasm is stressed.
Introduction The origin of the common bean (Phaseolus vulgaris L.) has been studied by several authors. Archaeological remains dated from 7000 B.C. were found in Tehuacfin, Mexico (Kaplan, 1965). Further evidences indicated that the common bean domestication is older in Andean America than in Mesoamerica. Therefore it was stated that domestication could have occurred independently in both regions or that the landraces and wild relative forms from Mesoamerica could have derived from South American ones (Kaplan, 1981). Recent studies, including electrophoretic analysis of the phaseolins, seed storage proteins, support the first hypothesis (Gepts & Bliss, 1985; Gepts, 1990). From the species of Phaseolus in Cuba, P. vulgaris is one of the most intensively cultivated crops
(Castifieiras & Shagarodsky, 1991; Esquivel et al., 1989). Variation of Cuban bean germplasm is not well known, since it is under-represented in mostly all bean germplasm collections (Bettencourt et al., 1989). Studies on the variation and origin of common bean material in America only included limited Cuban material (Voysest, 1983). In 1982 the INIFAT genebank started a program to collect plant genetic resources in Cuba, which was joined by IGK in 1986 (Hammer & Esquivel~ 1991). During collecting expeditions in Cuba, P. vulgaris was one of the species most often collected and a wide diversity has been detected for some seed characters (for a short review about collecting missions in Cuba see Esquivel et al., 1989). The knowledge of the genetic diversity available in a germplasm collection allows the breeders to choose the right strategy in the breeding program
m
scriptor list. The methodological principles proposed by Hernfindez Xolocotzi (1970) were followed to record additional ethnobotanical data such as vernacular names, origin of the landraces, uses,
Fig. 1. Graphic of the first (I), second (II) and third (III) principal components derived from Principal Component Analysis for agronomical characters.
(Lefort-Buson et al., 1988). Among the different mathematical methods used to study the variation within germplasm collections, multivariate analysis methods are included (Bedigian et al., 1986; Vanderborght, 1986; Hilling & Iezzoni, 1988; Peeters & Martinelli, 1989). Landraces have been considered as a valuable initial material for plant breeding, because they contain co-adapted gene complexes with tolerance or adaptation to diseases and specific ecological conditions respectively (Harlan, 1975). Accordingly landraces in Costa Rica showed tolerance to the web blight as a result of the selection pressure for that disease (Debouck et al., 1989). Considering the results of common bean collecting activities in Cuba and the possibility to study germplasm collections using multivariate statistical methods, the present study was done with the objectives to describe the variation existing in Cuban common bean germplasm, on the basis of a reduced number of important morpho-agronomical characters, to discuss the theories about the origin and dispersion of common bean at least in parts of America, and to demonstrate the use of valuable Cuban landraces to plant breeders.
Accessions from collecting missions were studied for two years at the experimental fields of INIFAT, and characterization/evaluation were carried out for the following minimum descriptor list proposed by Castifieiras (1990), which includes selected descriptors from the IBPGR list (IBPGR, 1982) and others in agreement with breeders' wishes. The following acronyms are used: 1. Cotyledons colour CC 2. Colour or trifoliate leaves CTL 3. Anthocyanin presence in the leaves AL 4. Growing habit GH* 5. Stem diameter (cm) SD* 6. Stem length (cm) SL* 7. Wings colour WC 8. Standard colour SC 9. Number of flowers per inflorescence NFI* 10. Pod form PF 11. Pod thickness (cm) PT 12. Pod width (cm) PW 13. Pod lenght (cm) PL 14. Pod colour at physiological maturity CPM 15. Pod cross section PCS 16. Pod beak form PBF 17. Dry pod consistency DPC* 18. Number of pods per plant NPP* 19. Number of seeds per pod NSP* 20. Seed form SF 21. Seed testa shine STS 22. Seed testa colour STC* Table 1. Distribution of bean accessions collected in Cuba according to size and grain colour Testa colour
Seed size
No. of accessions
Total %
black
small
176
53.7
red
small medium big
83 23 43
25.3 7.0 13,1
white
small medium
2 1
0.6 0.3
Material and methods
Collecting expeditions were carried out in Cuba during 1982-88. Passport information was gathered during expeditions, according to the IBPGR de-
3 23. Seed axis form SAF 24. Seed protein per cent SPP* 25. Weight of 100 seeds WHS* 26. Days to germination DG* 27. Days to flowering DF* 28. Days to physiological maturity DPM* 29. Flowering duration FD 30. Days to technical maturity DTM* 31. Hypocotyl pigmentation HP 32. Stem form SF* 33. Number of nodes per plant NNP* 34. Number of inflorescences per plant NIP* For statistical analysis descriptors were separated in agronomical (*) and botanical ones. Two methods were used: Principal Component Analysis (PCA) for agronomical data, because of prevailing quantitative characters, and Correspondence Factor Analysis (CFA) for botanical data (mostly qualitative characters). All statistical management was
Table 2. Distribution of 96 accessions of common bean according to important agronomic descriptors (for explanation see under Material and methods)
Descriptors
Descriptor states
Frequency %
GH
Type I Type II Type III 7-18 19-30 31-50 3-4 5-6 6-7 17.5-24.4 24.5-31.3 31.4-38.2 38.3-45.1 45.2-52.0 Erect Compact Irregular Runner 28-35 36-42 43-5O 71-83 84-97 98-111
3.12 84.24 12.48 38.48 57.20 4.16 5.20 61.36 33.28 87.36 7.28 2.08 1.04 2.08 69.68 21.84 7.28 1.04 15.60 82.16 2.O8 11.44 58.24 30.16
NPP
NSP
WHS
SF
DF
CPM
done using the statistical package STAT-ITCF, with an Olivetti M-24 microcomputer.
Results and discussion
Variation of common bean germplasm A total of 328 accessions of common bean was collected during the expeditions mentioned. Most of the material collected consists of traditionally cultivated landraces. In some areas the farmers maintain their old material, even so the main production is based on modern cultivars. The analysis of passport data shows that most of the people know and manage vernacular names related to seed testa colour, such as 'negro' (black), 'blanco' (white), 'rosado' (rosy), 'crema' (cream), 'bayo' (striped), 'mulatico' (brownish), etc. The vernacular name 'criollo' is frequently found for small black-seeded cultivars. Sometimes vernacular names are related to seed shape and form, as 'bollto' (rounded), 'largo' (large) and 'grande' (big). Names of regions are also given to bean landraces. Landraces with small seeds and black or red testa colour where most often found, few of them have white testa (Table 1). According to the distribution, red seeded landraces are more frequent in the Central provinces of Cuba, while the black ones predominate in the Western provinces. This generally agrees with the taste of people, but is also related to the tolerance to pests, diseases or unfavourable growing conditions. Black-seeded cultivats have a better taste and very few spices are needed to complement most Cuban dishes based on beans, which is the opposite as compared with the red and white-seeded cultivars. In traditional agriculture landraces are grown without watering and phytosanitary protection. Mostly red and black-seeded landraces are adapted to such conditions. The variation present can also be observed from descriptors appearing in Table 2. Even so most of the accessions belong to a single phenotype usually desirable for plant breeding, there are some of them in the extreme descriptor states. The results of PCA with agronomical characters
the importance of the first character for the classification of intraspecific variation. Figure 2 shows the possibility to differentiate groups according to the character Seed testa shine (STS), since group I is integrated with red accessions having shining testa, while group II includes black and white accessions, without shining testa. The intensity of purple colour of pods increases in accessions of group II, which differ from the accessions of group I in having lighter pod colour and few pods at the physiological maturity stage. ®
Origin of Cuban common bean germplasm Fig. 2. R e s u l t s o f t h e C o r r e s p o n d e n c e F a c t o r A n a l y s i s f o r b o tanical characters. Explanations of acronyms under Material and methods,
(Table 3) show that 58.8% of the total variation was accumulated in the three main components, which confirm the previous results of Castifieiras (1990) on the selection of a minimum descriptor list. Nevertheless, in the material studied there are characters with greater contribution, as Days to flowering (DF), Days to physiological maturity (DPM) and Seed testa colour (STC) to the first axis and Dry pod consistency (DPC), Number of inflorescences per plant (NIP) and Number of pods per plant (NPP) to the second axis. Figure 1 shows that Days to flowering (DF) and Days to physiological maturity (DPM) values increase to the right side of the C1 axis, so that three groups can be observed. Group I integrates early red seeded accessions, group II red intermediate accessions and group III brings together late black and white accessions. In the groups I and III prevail accessions with fibrous Dry pod consistency (DPC) and in group II accessions with silky and intermediate fibrous DPC. In the groups I and III accessions with erect Stem form (SF) dominate, whereas in group II accessions have erect, irregular or compact SF. The results of CFA with botanical characters (Table 4) allow to demonstrate that the characters Pod colour at physiological maturity (CPM) and Seed testa shine (STS) have a marked contribution to the two main factors, which allows us to confirm
The origin of the cultivated plants should be traced on the basis of the analysis of archaeological, historical, linguistic and botanical data (de Candolle, 1882), the archeological remains being the most important evidence. Due to the climatic conditions of Cuba, it has been nearly impossible to find remains of plants in the archeological sites of Indian Table 3. P r o p e r v a l u e s a n d v e c t o r s o f t h e a g r o n o m i c c h a r a c t e r s ( f o r e x p l a n a t i o n see u n d e r M a t e r i a l a n d m e t h o d s ) o f 96 b e a n accessions
Cl Variance
C3
C2
5.5352
1.5640
1.3800
% of contribution
32.6
18.0
8.2
cumulative %
32.6
50.6
58.8
Proper vectors
Coefficients
NRF
0.2821
0.1088
SL
0.2930
- 0.1838
0.3079
SD
0.1450
- 0.2041
- 0.4272
NNP
0.2998
0.2028
0.0273
SF
0.1546
0.0518
0.4683
GH
0.2715
- 0.1799
0.3597
NIP
0.2505
0.3870
- 0.0850
NPP
0.2706
NSP
0.2705
WHS
0.3570 -0.0236
- 0.1246
- 0.1649 -0.1119
- 0.2748
- 0.0749
0,0674
STC
0.3176
- 0.1972
- 0.1677
DPC
0.0099
- 0.5766
- 0.0996
DG
0.1033
- 0.2958
- 0.1193
DF
0.3610
- 0.0790
- 0.0284
DPM
0.3372
- 0.1902
0.0305
DTM
- 0.0418
0.1678
0.2030
0.0542
- 0.1540
- 0.4598
SPP
cultures (Guarch, pers. comm.), therefore the indirect evidences from other sources become more important. The origin of the common bean germplasm in Cuba should be analyzed in space and time, i.e. this problem could be divided in the following questions: from where, when and how did the common bean arrive to Cuba? The existence of two genepools resulting from independent domestications in Mesoamerica and in the southern Andes has recently been established for the common bean (Gepts, 1988; Gepts et al., 1986). A correspondence between seed size and phaseolin type was found for each centre. Beans with small seeds and phaseolin type S were domesticated in Mesoamerica, while beans with larger seeds and phaseolin type T originated in the Andean centre (Gepts & Bliss, 1986). The results of this paper show that there are two main groups of landraces in Cuba according to seed size (Table 2), 87% of the accessions have Weights
Table 4. Proper values and absolute contribution of botanical descriptors (for explanation see under Material and methods) for 96 bean accessions
Proper values % of contribution cumulative %
C~
Cz
0.0137 42.9 42.9
0.0043 13.9 56.8
% of contribution of original variables CC HP CTL AL SC WC PF SAF STS SF CPM PCS PL PW GV PBF FD
2.2 0.1 0.4 0.1 3.2 0.9 0.6 1.4 23.1 9.7 29.0 0.2 6.9 0.7 0.6 0.8 0.8
1.2 0.0 0.0 1.6 0.7 0.0 0.1 1.0 12.2 5.7 59.6 0.0 0.3 0.0 0.0 2.0 2.3
Fig. 3. Ways of introduction of Phaseolus vulgaris to Cuba. T = Pre-Columbian introduction of T phaseolin types. S = PostColumbian introduction of S phaseolin types.
of 100 seeds (WHS) between 17.5-24.4 g, whereas the rest has larger seeds with WHS up to 45.2 g. This agrees with the results of Lioi et al. (1990) who studied the phaseolin types of 119 accessions of common bean collected in Cuba, including the material referred to in this paper. 62% of the accessions showed phaseolin type S and 38% a T phaseolin type. The common S phaseolin type is associated with small black or red seeded landraces, with average WHS of 21 g, whereas the T phaseolin type comprises mostly large-seeded landraces with pink, red or mottled red seeds and average WHS of 37.5g. The predominance of landraces with the phaseolin type S and the absence of the phaseolin type 'C', resemble the pattern already found in other Caribbean Islands by Gepts et al. (1988), when Cuban material was not included. It is evident that common bean germplasm in Cuba has been introduced from both major domes-
tication centres. Voysest (1983) discussed the origin of common bean varieties in Latin America. He proposed the same dispersal routes as Mackie (1943) for the different cultigroups of P. lunatus. He suggested that the small-seeded common bean cultivars of the Mexican coast, Central America, the Caribbean Islands, Venezuela and Brazil have a common origin. Esquivel et al. (1990) already discussed the probable migratory routes of P. lunatus in the Caribbean on the basis of morphological data from wild, weedy and cultivated material recently collected in Cuba. Lioi et al. (1991) also reported the presence of the Mesoamerican and Andean phaseolin types for lima bean in Cuba. These results, together with the actual knowledge about the socioeconomical relations in pre- and post-Columbian times in Mesoamerica and the Caribbean, gave no support to Mackie's hypothesis. One of the main preconditions on which Mackie's hypothesis is based, is the existence of pre-Columbian commercial contacts between Mexico and Cuba. Dacal and Rivero de la Calle (1984) give a detailed information about the first inhabitants of Cuba. Geographical and archaeological evidences indicate that such contacts were not possible before Columbus. On the other hand, historical data show that beans were already in Cuba at the arrival of the Europeans. In the navigation diary of Columbus (Col6n, 1493) he reports beans different from those existing in Spain (Vigna unguiculata and Vicia faba). Probably the first introductions to Cuba were the large-seeded cultivars with phaseolin type T, which could have arrived with the 'Tainos', Arawak Indian tribes who left the Eastern slopes of the Andes and reached Cuba about 950 B.C. (Guarch, 1978; Dacal & Rivero de la Calle, 1986). Cultivars with small seeds and phaseolin type S could have arrived via the first route proposed by Gepts et al. (1988) starting from Mexico and following the Caribbean coast to Colombia, Venezuela and Brazil, and then by the same route through the Antilles. It is also possible that both types were later on reintroduced. After the Conquest true commercial routes which connected Cuba with continental America were established, because it was the con-
fluence place of the fleet which returned to Spain with merchandise from all American colonies, which notably contributed to the transculturation process. Linguistic evidences demonstrate the influence of different Indoamerican cultures. 42% of Indoamerican words in the Spanish spoken in Cuba are derived from the insular Arawak (Taino language), but the rest from other languages, as Carib, Nahualt, Maya, Tupi-Guarani and even Quechua (Vald6s 1978, 1986). Accordingly, the S type seeds were possibly reintroduced directly from Mexico after the Conquest, and T types from Peru via Panama. Figure 3 shows the possible routes of introduction of common beans. As stressed by Sauer (1962) the agriculture of Central America was the agriculture of grains, based on maize-bean cultivation. Kaplan (1965) considered that the archaeological studies of Phaseolus beans have been linked with those of maize. That is a clear indication of the importance of a combined role of these crops in the life of ancient populations. So Gepts et al. (1988) considered that the dissemination routes for beans show some parallels with those reported by McClintock et al. (1981) for maize.
Utilization of common bean germplasm According to the results of preliminary evaluation, landraces with agreeable characteristics could be selected and therefore many of them have been incorporated as genetic stocks in the bean breeding program of INIFAT. Table 5 provides a summary
Table5. Resultsof the yieldevaluation(kg/ha)of commonbean landraces Seed colour
Landraces
Commercialvarieties
Name
yield
Name
Black
P242-1
1759
GOira 89 1686
Red
P186 P219
1640 1634
Rosas
White
P500 P384
1531 1408
Bonita 11 1366
yield
1356
7 of bean yield evaluation trials under low farming inputs, where some landraces are compared with commercial varieties, according to Castifieiras (1988, 1990). Further studies of red type landraces in four localities under production conditions revealed yields of 2610 kg/ha and 2420 kg/ha for P186 and P219 respectively, while the commercial variety Rosas had a yield of 1956 kg/ha (Castifieiras et al., 1989). Several bean landraces showed also better resistance against Bean Common Mosaic Virus (BCMV) and rust (Guerra y Lastres, 1987; Guerra et al., 1990). Additional studies on the resistance against the former diseases demonstrated that P186 showed resistance to NL-7 and NL-1 races of BMCV and rust races 5, 8 and 10, while P219 showed resistance to the same BMCV races and to rust races 5, 7, 8 and 10 (Castifieiras et al., 1989).
Acknowledgement The authors dedicate this paper to the memory of the great Mexican ethnobotanist Efraim Hermindez Xolocotzi who died last February. He devoted his life to the study of plant genetic resources in Latin America.
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