Lathyrus improvement for resistance against biotic and abiotic stresses: From classical breeding to marker assisted selection

Summary Several Lathyrus species and in particular Lathyrus sativus (grass pea) have great agronomic potential as grain and forage legume, especially in drought conditions. Grass pea is rightly considered as one of the most promising sources of calories and protein for the vast and expanding populations of drought-prone and marginal areas of Asia and Africa. It is virtually the only species that can yield high protein food and feed under these conditions. It is superior in yield, protein value, nitrogen fixation, and drought, flood and salinity tolerance than other legume crops. Lathyrus species have a considerable potential in crop rotation, improving soil physical conditions; reducing the amount of disease and weed populations, with the overall reduction of production costs. Grass pea was already in use in Neolithic times, and presently is considered as a model crop for sustainable agriculture. As a result of the little breeding effort invested in it compared to other legumes, grass pea cultivation has shown a regressive pattern in many areas in recent decades. This is due to variable yield caused by sensitivity to diseases and stress factors and above all, to the presence of the neurotoxin β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP), increasing the danger of genetic erosion. However, both L. sativus and L. cicera are gaining interest as grain legume crops in Mediterranean-type environments and production is increasing in Ethiopia, China, Australia and several European countries. This paper reviews research work on Lathyrus breeding focusing mainly on biotic and abiotic resistance improvement, and lists current developments in biotechnologies to identify challenges for Lathyrus improvement in the future.     

Faba bean breeding for resistance against biotic stresses: Towards application of marker technology

Summary Faba beans are adversely affected by numerous fungal diseases leading to a steady reduction in the cultivated area in many countries. Major diseases such as Ascochyta blight (Ascochyta fabae), rust (Uromyces viciae-fabae), chocolate spot (Botrytis fabae), downy mildew (Peornospora viciae) and foot rots (Fusarium spp.) are considered to be the major constraints to the crop. Importantly, broomrape (Orobanche crenata), a very aggressive parasitic angiosperm, is the most damaging and widespread enemy along the Mediterranean basin and Northern Africa. Recent mapping studies have allowed the identification of genes and QTLs controlling resistance to some of these diseases. In case of broomrape, 3 QTLs explained more than 70% of the phenotypic variance of the trait. Concerning Ascochyta, two QTLs located in chromosomes 2 and 3 explained 45% of variation. A second population sharing the susceptible parental line also revealed two QTLs, one of them likely sharing chromosomal location and jointly contributing with a similar percentage of the total phenotypic variance. Finally, several RAPD markers linked to a gene determining hypersensitive resistance to race 1 of the rust fungus U. viciae-fabae have also been reported. The aim of this paper is to review the state of the art of gene technology for genetic improvement of faba bean against several important biotic stresses. Special emphasis is given on the application of marker technology, and Quantitative Trait Loci (QTL) analysis for Marker-Assisted Selection (MAS) in the species. Finally, the potential use of genomic tools to facilitate breeding in the species is discussed. The combined approach should expedite the future development of lines and cultivars with multiple disease resistance, one of the top priorities in faba bean research programs.     

Current and future strategies in breeding lentil for resistance to biotic and abiotic stresses

Lentil production is limited by lack of moisture and unfavorable temperatures throughout its distribution. Waterlogging and salinity are only locally important. Progress has been made in breeding for tolerance to drought through selection for an appropriate phenology and increased water use efficiency and in breeding for winter hardiness through selection for cold tolerance.The diseases rust, vascular wilt, and Ascochyta blight, caused by Uromyces viciae-fabae, Fusarium oxysporum f. sp. lentis, and Ascochyta fabae f. sp. lentis, respectively, are the key fungal pathogens of lentil. Cultivars with resistance to rust and Ascochyta blight have been released in several countries and resistant sources to vascular wilt are being exploited. Sources of resistance to several other fungal and viral diseases of regional importance are known. In contrast, although the pea leaf weevil (Sitona spp.) and the parasitic weed broomrape (Orobanche spp.), and to a lesser extent the cyst nematode (Heterodera ciceri), are significant yield reducers of lentil, no sources of resistance to these biotic stresses have been found. Directions for future research in lentil on both biotic and abiotic stresses are discussed.     

Current status and future strategy in breeding chickpea for resistance to biotic and abiotic stresses

Chickpea (Cicer arietinum L.) production has remained static for the past two decades. One major limiting factor has been susceptibility of cultivars to several biotic and abiotic stresses that adversely affect yield. In recent years, cultivars resistant to Ascochyta blight (Ascochyta rabiei [Pass.] Lab.), Fusarium wilt (Fusarium oxysporum f. sp. ciceris), and cold have been bred and released in many countries. Some progress has been made in breeding for resistance to drought, insects, and cyst nematode, but not for viruses, heat, and salinity. Two or more stresses are of equal importance in most chickpea growing areas. Therefore, future efforts should be directed toward the development of cultivars with multiple-stress resistance. Proper understanding of important stresses in different countries and the genetics of resistance should lead to more systematic approaches to resistance breeding. Wild Cicer species hold promise and deserve attention in resistance breeding.     

Current status and future strategy in breeding pea to improve resistance to biotic and abiotic stresses

The economic importance and current progress made in studies of the host-parasite relationship and identification of sources of resistance and breeding strategies of some important biotic diseases of pea are reviewed in this paper. The root rot complex caused by Rhizoctonia solani, Fusarium solani, Aphanomyces euteiches, Pythium ultimum and Fusarium oxysporum f. sp. pisi, race 1 and 2 has been reported from all commercial pea growing areas of the world. Adequate sources of resistance have been identified and there has been impressive success in the control of the Fusarium wilt pathogen following the introduction of wilt-resistant cultivars. Leaf and stem diseases of pea caused by the Ascochyta complex, Peronospora viciae and Erysiphe pisi are prevalent in most temperate pea growing regions of the world. Several sources of resistance are available, some of which are surprisingly durable. The biochemical genetic parameters of phenolic content used for assaying resistance to Erysiphe pisi offers an alternative method of evaluating breeding material. Wild relatives of pea (Pisum fulvum and P. humile) are valuable additional sources of genetic variation and provide good sources of resistance to pests and diseases. In temperate rainfed pea growing areas of southern Australia, pea seed yield is more closely related to dry matter production than harvest index. Tall and leafy cultivars proved more productive than afila types.     

Present status and future strategy in breeding faba beans (Vicia Faba L.) for resistance to biotic and abiotic stresses

Progress is being made, mainly by ICARDA but also elsewhere, in breeding for resistance to Botrytis, AScochyta, Uromyces, and Orobanche; and some lines have resistance to more than one pathogen. The strategy is to extend multiple resistance but also to seek new and durable forms of resistance. Internationally coordinated programs are needed to maintain the momentum of this work.Tolerance of abiotic stresses leads to types suited to dry or cold environments rather than broad adaptability, but in this cross-pollinated species, the more hybrid vigor expressed by a cultivar, the more it is likely to tolerate various stresses.     

Pyramiding genes for resistance to bean common mosaic virus

Summary Gene pyramiding in Phaseolus vulgaris is being utilized to develop more effective resistance to the temperature-insensitive-necrosis-inducing (TINI) strains of Bean Common Mosaic Virus (BCMV) present in the USA. Our data indicate that contrary to previous work, the bc-3 gene is effective against these strains in the absence of the strain unspecific bc-u gene in genotypes possessing the I gene. The epistatic bc-3 gene interferes with traditional efforts to pyramid the other recessive resistance genes by masking their activity. Indirect selection based on markers linked to the other recessive resistance genes would likewise be ineffective without the ability to also select for the bc-u gene which is required for expression of the bc-2 ² gene in germplasm carrying the I gene. Because the most resistant genotype (I, bc-u, bc-I ², bc-2², bc-3) can only be introduced into a wide range of germplasm through the use of molecular markers linked to the different resistance genes, the search for a marker linked to the strain unspecific bc-u gene should also be given priority.     

Introgressing resistance to bacterial and viral diseases from the middle American to Andean common bean

The genetic base of cultivars within market classes of common bean (Phaseolus vulgaris L.) is narrow. Moreover, small- and medium-seeded Middle American cultivars often possess higher yield and resistance to abiotic and biotic stresses than their large-seeded Andean counterparts. Thus, for broadening the genetic base and breeding for higher yielding multiple stress resistant Andean cultivars use of inter-gene pool populations is essential. Our objective was to determine the feasibility of introgressing resistance to Been common mosaic virus (BCMV, a potyvirus), and the common [caused by Xanthomonas campestris pv. phaseoli (Xcp) and X. campestris pv. phaseoli var. fuscans (Xcpf)] and halo [caused by Pseudomonas syringae pv. phaseolicola (Psp)] bacterial blights from the Middle American to Andean bean, using gamete selection. Also, we investigated the relative importance of the use of a landrace cultivar versus elite breeding line as the last parent making maximum genetic contribution in multiple-parent inter-gene pool crosses for breeding for resistance to diseases. Two multiple-parent crosses, namely ZARA I = Wilkinson 2 /// ‘ICA Tundama’ / ‘Edmund’ // VAX 3 / PVA 773 and ZARA II = ‘Moradillo’ /// ICA Tundama / Edmund // VAX 3 / PVA 773 were made. From the F₁ to F₅ single plant selection was practiced for resistance to the common and halo bacterial blights in both populations at Valladolid, Spain. The parents and F₅-derived F₆ breeding lines were evaluated separately for BCMV, and common and halo bacterial blights in the greenhouse at Filer and Kimberly, Idaho in 2001. They were also evaluated for the two bacterial blights, growth habit, seed color and 100-seed weight at Valladolid in 2002. All 20 F₁ plants of ZARA I were resistant or intermediate to common and halo bacterial blights in the greenhouse, but their F₂ and subsequent families segregated for both bacterial blights. Segregation for resistant, intermediate, and susceptible plants for common bacterial blight occurred in the F₁ of ZARA II. Simple correlation coefficient for common bacterial blight between the F₁ and F₁-derived F₂ families was positive (r = 0.54 P < 0.05) for ZARA II. From the F₂ to F₅ the number of families resistant to both bacterial blights decreased in both populations. Only four of 20 F₁ plants in ZARA I resulted in seven F₆ breeding lines, and only one of 32 F₁ plants in ZARA II resulted in one F₆ breeding line resistant to the three diseases. None of the selected breeding lines had seed size as large as the largest Andean parent. The use of elite breeding line or cultivar as the last parent making maximum genetic contribution to the multiple-parent inter-gene pool crosses, relatively large population size in the F₁, and simultaneous selection for plant type, seed traits as well as resistance to diseases would be crucial for introgression and pyramiding of favorable alleles and quantitative trait loci (QTL) of interest between the Andean and Middle American beans.     

Gamete selection for improving physiological resistance to white mold in common bean

White mold (WM), caused by Sclerotinia sclerotiorum (Lib.) de Bary, is a widespread disease of dry and green bean (Phaseolus vulgaris L.) in North America. Gamete selection (GS) was effective to combine and pyramide resistant genes and quantitative trait loci (QTL) for common bacterial blight. Our objective was to determine the effectiveness of GS to introgress physiological resistance to white mold. Two inter-gene-pool double-cross populations were developed. Selection for WM resistance was practiced from F₁ to F₄. Thirteen selected F_1:5 breeding lines of each population and their four parents were evaluated. Two separate inoculations were made on each plant 1 week apart using a cut-stem method. The WM reaction was scored at 16, 23, and 33 days post inoculation (DPI) using a scale from 1 (no disease) to 9 (severely diseased or dead). In F₁, 52% of Pop I (USPT-WM-1/CORN 601//USPT-CBB-1/92BG-7) and 67% of Pop II (Chase/I9365-25//ABL 15/A 195) susceptible plants were discarded. In F_4, only 1.2% of families from Pop I, and 0.9% for Pop II, survived the selection process. An average of 20.5% gain in WM resistance was obtained for both populations in F₄. Four breeding lines of Pop I had significantly (P = 0.05) lower WM score (4.1–4.6) and four were equal (4.7–4.9) to the best WM-resistant parent 92BG-7 (4.9), while ten breeding lines of Pop II were equal (4.5–4.8) to the best WM-resistant parent A 195 (4.6). Thus, GS was effective for improving WM resistance in common bean.     

Molecular markers and allelic relationships of anthracnose resistance gene cluster B4 in common bean

Angular leaf spot is one of the major diseases of the common bean. The extensive genetic variability of this pathogen requires the constant development of new resistant cultivars. Different sources of resistance have been identified and characterized. For the State of Minas Gerais, Brazil, four main resistance sources were found: Mexico 54, AND 277, MAR 2 and Cornell 49-242. Independent characterization of these genotypes demonstrates that resistance in all four sources is dominant and monogenic. However, there are no studies on the relationship and independence of these genes. In the present work, allelism tests were carried out to understand the relationship among the resistance genes present in these four resistance sources. The data revealed a much higher complexity in the resistance inheritance of these genes than previously reported. It was demonstrated that Cornell 49-242 possesses a dominant gene (Phg-3); Mexico 54 possesses three genes, denominated Phg-2, Phg-5 and Phg-6. In MAR 2, two genes were found, one independent designated Phg-4 and the other, an allelic form of Phg-5, denominated of Phg-5². Allelic forms were also found in AND 277, Phg-2², Phg-3² and Phg-4². These results have special importance for breeding programs aiming to pyramid resistance genes.     

An allelic series at the Co-1 locus conditioning resistance to anthracnose in common bean of Andean origin

In this study, we characterized the genetic resistance of the Andean bean cultivars Kaboon and Perry Marrow and their relation to other sources of anthracnose resistance in common bean. Based on the segregation ratio (3R:1S) observed in two F₂ populations we demonstrated that Kaboon carries one major dominant gene conferring resistance to races 7 and 73 of Colletotrichum lindemuthianum. This gene in Kaboon is independent from the Co-2 gene and is an allele of the Co-1 gene present in Michigan Dark Red Kidney (MDRK) cultivar. Therefore, we propose the symbol CO-1 ² for the major dominant gene in Kaboon. The Co-1 is the only gene of Andean origin among the Co anthracnose resistance genes characterized in common bean. When inoculated with the less virulent Andean race 5, the segregation ratio in the F₂ progeny of Cardinal and Kaboon was 57R:7S (p = 0.38). These data indicate that Kaboon must possess other weaker dominant resistance genes with a complementary mode of action, since Cardinal is not known to possess genes for anthracnose resistance. Perry Marrow, a second Andean cultivar with resistance to a different group of races, was shown to possess another resistant allele at the Co-1 locus and the gene symbol Co-1 ³ was assigned. In R × R crosses between Perry Marrow and MDRK or Kaboon, no susceptible F₂ plants were found when inoculated with race 73. These findings support the presence of a multiple allelic series at the Andean Co-1 locus, and have major implications in breeding for durable anthracnose resistance in common bean. This revised version was published online in July 2006 with corrections to the Cover Date.     

Breeding the common bean (Phaseolus vulgaris L.) for resistance to bean common mosaic virus: alternatives to backcrossing

A series of field experiments was undertaken in order to determine whether resistance to bean common mosaic virus (BCMV) could be incorporated into genotypes of the common bean (Phaseolus vulgaris L.) suitable for cultivation in Zimbabwe without recourse to backcrossing. Six inbred genotypes carrying the resistance-conferring alleles at the loci I and Bc-3 were crossed with five locally-adapted inbred genotypes. The first experiment comprised F₃ progeny rows, each derived from a single unselected F₂ plant, the second, F₃ bulks selected for resistance, and the third, a comparison of selected and unselected F₂-derived F₄ lines. The number of days to flowering and to maturity, the incidence of mosaic and necrosis symptoms, seed yield and seed size were recorded. There was evidence that late flowering and maturity were associated with BCMV resistance in some crosses, though not strongly enough to present an obstacle to plant breeding. The incidence of virus symptoms and seed yield were influenced by genetic factors additional to the major resistance genes, and variation in seed yield was present not only between bulk populations of crosses, but also between single-row plots of lines within crosses. This indicates that early-generation selection for yield in the presence of BCMV, even among progeny selected for BCMV-resistace, is likely to be effective. However, the variation in yield among F₄ lines was least in the highest-yielding crosses, which may represent a limit to successful selection for yield. Seed size was partly under additive genetic control, but there was also evidence of non-allelic interactions. There was no association between large seed size, preferred by consumers, and susceptibility to BCMV in the progeny, indicating that the association between these characters in the parent lines is fortuitous and will not present an obstacle to plant breeding. It is noted that a considerable amount of useful genetic information can be obtained without recourse to elaborate crossing schemes, provided that unselected progeny are included in experiments as controls. The evidence presented indicates that resistance to BCMV can be combined with appropriate values of maturity date, yield and seed size without the need for backcrossing.     

Use of molecular markers to accelerate the breeding of common bean lines resistant to rust and anthracnose

The main goal of this work was to introduce resistance genes for rust, caused by Uromyces appendiculatus, and anthracnose, caused by Colletotrichum lindemuthianum, in an adapted common bean cultivar through marker-assisted backcrossing. DNA fingerprinting was used to select plants genetically closer to the recurrent parent which were also resistant to rust and to race 89 of C. lindemuthianum. DNA samples extracted from the resistant parent (cv. Ouro Negro), the recurrent parent (cv. Rudá), and from BC₁, BC₂ and BC₃ resistant plants were amplified by the RAPD technique. The relative genetic distances in relation to the recurrent parent varied between 9 and 59% for BC₁, 7 and 33% for BC₂, and 0 and 7% for BC₃ resistant plants. After only three backcrosses, five lines resistant to rust and anthracnose with, approximately, 0% genetic distance in relation to the recurrent parent were obtained. These lines underwent field yield tests in two consecutive growing seasons and three of them presented a good yield performance, surpassing in that sense their parents and most of the reference cultivars tested.     

Evaluation of resistance sources and inheritance of resistance in kidney bean to Indian virulences of Colletotrichum lindemuthianum

Summary Forty nine common bean lines comprising of exotic accessions and locally grown cultivars evaluated against Colletotrichum lindemuthianum exhibited differential resistance to its races in Himachal Pradesh, a north-western Himalayan state of India. Some exotic accessions like G 2333, Cornell 49242, PI 207262, Mexique 222, TO, Perry Marrow, Kaboon and Widusa were resistant to more than five Indian races, whereas two Indian accessions KRC-5 and Hans showed resistance to six and four races, respectively. However, nine accessions KRC-8, KR-40, KR-43, KR-81, KR-62-2, KR-90, KR-142, KR-148, and KR-216 were resistant to three races. Race specific resistance has been observed in different bean cultivars. Studies on inheritance of resistance in exotic accession G 2333 and Indian accession, KRC-5 showed that two independent dominant genes conferred resistance in G 2333 to race 3 and 515 and a single dominant gene controlled resistance in KRC-5 to race 775, indicating resistance from these sources is easily transferable to the locally adapted susceptible cultivars.     

Inheritance of resistance to Acanthoscelides obtectus in a wild common bean accession crossed to commercial bean cultivars

Summary The bean weevil Acanthoscelides obtectus, is an important storage pest of common beans in Latin America and Africa. A few wild bean accessions from Mexico have been identified as highly resistant to the weevil. One accession, G 12952, was crossed to two susceptible bean cultivars differing in seed size. Reciprocal F1 and F2 individual seed were evaluated for days to adult emergence (DAE) and emerged adult weight. Maternally inherited seed size affected resistance measurements only in the F1 reciprocal crosses, however, the overall resistance level of the F1 was more similar to that of the susceptible cultivars. The F2 showed a continuous, but skewed distribution from low to high DAE. Very few F2 individuals had the resistance level of G 12952. When the frequency distributions were divided into discrete categories based on parental response, resistance was found to be inherited as two recessive complementary genes. The F3 generation showed an overall lowering of resistance levels compared to their original F2 evaluations. However, none of the lines classified as resistant (50 DAE) in the F2, fell into the susceptible category in the F3, indicating that the resistant genotypes were relatively stable as expected with recessively inherited traits. Modifying genes from the commercial parents may be responsible for general lowering of resistance. Seed size was negatively correlated with adult weight but not with DAE. The unique resistance of the wild bean accessions is discussed in relation to its inheritance. The results and obstacles encountered in the A. obtectus breeding program at CIAT are described.     

Linkage of dominant hypersensitive resistance to bean common mosaic virus to seed color in Phaseolus vulgaris L.

Summary Evaluation of Phaseolus vulgaris germplasm bank materials and progenies from a large number of crosses using red- or yellow-colored, BCMV-susceptible bean lines, crossed to purple- or grey/brown-colored, hypersensitive-resistant lines, suggested strong trait association between seed color and BCMV resistance. The cross of red-mottled I⁺I⁺ (susceptible) BAT 1255R to isogenic purple-mottled II (resistant) BAT 1255M was made to study the segregation of the two characters and to recover red-mottled resistant progenies. No recombinant genotypes were observed among 353 F₃ families inoculated with BCMV-NL3, suggesting that linkage of purple-mottled seed color and dominant BCMV resistance is very close.Contribution of the Centro Internacional de Agricultural Tropical.     

Development of an appropriate breeding scheme for tolerance to Empoasca kraemeri in common bean

Summary Significant advances in increasing tolerance to the leafhopper Empoasca kraemeri Ross & Moore in common bean have been obtained using a new breeding scheme where yield under leafhopper attack is the principal selection criterion in the evaluation of progenies. However, to further refine this breeding scheme, a study was conducted to determine whether selection for nonprotected yield would be more effective in early versus late generations. Two selection strategies were compared. In Strategy I, early generation selections in the F₂ and F₃ were compared to Strategy II where late generation selections were made in the F₄ and F₅, with the F₂ and F₃ generations advanced using single pod descent and bulk practices, respectively. Yield trials of the F₆ lines from both selection strategies were conducted under nonprotected and insecticide protected treatments. No significant differences were detected between the two selection strategies. However, Strategy II did produce advanced lines with greater nonprotected yields than did Strategy I, with the best F₆ line, in three out of four crosses, coming out of Strategy II. Late generation selection is recommended over early generation selection. Results of the new leafhopper breeding scheme, based on yield, are compared to the old breeding scheme where selections were made using visual selection practices in early generations.     

Development of common bean (Phaseolus vulgaris L.) lines resistant to the bean pod weevil,Apion godmani Wagner,in Central America

Summary The larva of the bean pod weevil (BPW), Apion godmani Wagner (Coleoptera: Curculionidae), causes serious yield losses in common bean (Phaseolus vulgaris L.) in Mexico and Central America, by consuming the seed as it develops in the immature pod. Resistance to the BPW was identified in bean germplasm of highland Mexican origin, and these sources of resistance were incorporated into a pedigree breeding program to recover locally adapted lines resistant to Bean Common Mosaic Virus and BPW, with commercial grain for Guatemala, Honduras and El Salvador. These lines yielded as well as or better than local cultivars in the absence of the insect, and better than local cultivars when the BPW was present. Resistance appeared to be governed by several genes, and was stable across geographic areas, seasons and planting systems.     

Breeding methods and selection indices for improved tolerance to biotic and abiotic stresses in cool season food legumes

The objective of breeding for stress tolerance is to improve productivity for a target level of stress. If tolerance is viewed as resistance to change in productivity with increasing stress, productivity under stress depends not only on stress tolerance, but also on maximum productivity. Index selection theory indicates that selection in non-stress environments will be more effective than direct selection for productivity under stress whenever the correlation between the two types of environments exceeds the heritability of productivity under stress. With high genetic correlation, selection should be conducted within a level of stress that maximizes heritability. In cases where heritability under non-stress is much higher than under stress, an index combining data from stress and non-stress environments is expected to be more efficient than selection based on evaluation only within stress environments.Secondary traits will be useful in breeding for productivity under stress whenever they have high heritability and high genetic correlation with productivity under stress. For some abiotic stresses and many biotic stresses, heritability will be highest in the presence of stress and indirect or index selection will be of limited value.     

Breeding for resistance to bean golden mosaic virus in an interracial population of Phaseolus vulgaris L.

Summary Eighty-three F₂-derived F₈ recombinant inbred lines (RILs) randomly selected from a population (TY 8419) of two common bean genotypes (Pinto UI 114 x ICA Pijao), the two parents, and a control cultivar (Topcrop) were screened for their reaction to bean golden mosaic virus (BGMV) under glasshouse conditions. A randomized complete block design with two replications was used. Data were recorded for disease incidence, foliar yellowing, number of pods and seeds, and yield.Significant (P<0.05) differences among the 83 RILs were recorded for all traits. Of these RILs, 11 lines did not develop any symptom, 24 lines had a mean disease incidence of 8%, 28 lines had a disease incidence of 26.6% and developed intermediate mosaic symptoms, and 20 lines were more susceptible (>50% disease incidence and severe yellowing) than either of the parents, which had 25% to 38% disease incidence and moderate symptoms. Heritability for disease incidence was 0.54±0.01, for foliar yellowing 0.36±0.13, for pods plant^-1 0.69±0.01, for seeds plant^-1 0.54±0.09, and for yield plant^-1 0.49±0.01. Gains from selection (at 20% selection pressure) for these traits were 49.5%, 31.6%, 24%, 0%, and 20%, respectively, over the mean value of all RILs. Correlation coefficients between disease incidence and foliar yellowing and among yield and its components were positive. Disease incidence and foliar yellowing showed negative associations with yield and its components.