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.     

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.     

Selection for seed yield in Andean intra-gene pool and Andean × Middle American inter-gene pool populations of common bean

Yields of large-seeded Andean (A) common bean (Phaseolus vulgaris L.) cultivars of Chile and Nueva Granada races are 40 to 60% lower compared to their Middle American (M) counterparts of small-seeded Mesoamerica and medium-seeded Durango races. Our objective was to use the concept of congruity backcrossing between Andean x Middle American inter-gene pool [AM 11833 = A 483 (A)///// A 686 (M) //// PVA 800A(A)/// ‘Carioca’ (M)// Carioca (M)/ G 19833 (A)] and between races within Andean gene pool [i.e., intra-gene pool Andean, AA 11834 = A483 (A)//// ‘Cardinal’(A) /// ‘Blanco Español’(A) // BlancoEspañol (A) / ‘Taylor’ (A)] to compare selection for seed yield improvement of large-seeded Andean beans. Seven hundred sixty seven F₂-derived F₃ (F_2:3) families were produced for each population. Visual appraisal for total plant performance, combined with seed yield from non-replicated plots was used for selection of 551 families in F_2:3, 182 families in F_2:4, and 91 families in F_2:5 in each population. Eight hundred twenty three F_5:6 lines were developed from the 91 F_2:5 families in each population. Visual selection, combined with seed yield in non-replicated plots was again used to select 294 lines in F_5:6 in each population. Similarly, 44 highest yielding F_5:7 lines were selected in AM 11833 and 39 F_5:7 lines in AA 11834. Thus, single plant selections were made in the F₂ and F₅, and plants within each plot were harvested in bulk in F₃, F₄, F₆, and F₇. Thirty nine F_5:8 lines from AA 11834 and 44 lines from AM 11833, parents, and checks were evaluated at Popayán and Quilichao, Colombia in 1998 and 1999. Selected lines in both populations, on average, out-yielded the mean of their large-seeded Andean parents. Mean yield of the lines selected from AM 11833 was 50% higher than AA 11834 lines. Twelve F_5:8 lines out-yielded the highest yielding Andean parents G 19833 and A 483 in AM 11833, whereas only one line yielded significantly higher (p < 0.05) than the highest yielding parent A 483 in AA 11834. However, none of selected lines out-yielded small-seeded Middle American parents used in AM 11833 (A 686 and ‘Carioca’). The mean 100 seed-weight of AA 11834 was 36 g compared to 28 G for AM 11833 F_5:8 lines. Selected lines had similar days to maturity as parents in AM 11833, and matured 3 d later in AA11834. Correlation coefficients between yield and 100 seed-weight were negative in both populations. Yield and days to maturity were positively correlated in AA 11834.     

Phenotypic recurrent selection in common bean (Phaseolus vulgaris L.) based on performance of S2 progenies

Summary The effect of a recurrent selection procedure for improving seed yield per plant was evaluated in common bean. Progress was registered after three cycles of recurrent selection with S2 progeny evaluation performed on the grain yield/plant. Gain from selection was estimated by field testing 45 random S2 lines from each of the C0, C1 and C2 populations. Response to selection for seed yield/plant was 6 g from C0 to C1 and 4.4 g from C1 to C2. The seed yield improvement was due principally to an increase in pods/plant; little contribution was provided by the 1000-seed weight and seeds/pod traits. Broad-sense heritability and genotypic variance for grain yield remained high in all cycles which suggests further gain from additional selection cycles. Grain yield/plant is significantly correlated to pods/plant and seeds/pod traits in all cycles, indicating that plants with favourable combinations of these traits could be identified. The proposed recurrent selection procedure appears to be effective in improving the population and extracting superior genotypes for varietal development.     

Sources and inheritance of earliness in tropically adapted indeterminate common bean

Summary Nineteen early maturing, indeterminate common bean parents were crossed with a tester line of normal maturity. All parents, F₁ hybrids, F₂'s, and backcross generations were evaluated at CIAT-Palmira, Colombia. All 19F₁ populations flowered and matured almost as early as the respective early parent (mean of 61 vs 59 days). The F₂ populations were slightly less early (63 days), but were still earlier than the mean of the two parents (65 days). Additive effects for days to maturity were usually larger than dominance or interaction effects. Narrow sense heritability for days to flower and maturity were high (greater than 0.67), while heritabilities of absolute and relative duration of pod filling were low (0.17 to 0.50). Also, our data indicated that for each day of increase in earliness, yield potential will decrease 74 kg/ha.     

Genetic progress after four cycles of recurrent selection for yield and grain traits in common bean

The objective of this study was to evaluate the genetic progress after four cycles of recurrent selection in common bean. The base segregating population was obtained from 10 parents, and derived the S_0:1 and S_0:2 families that were evaluated. The S_0:3 families with higher grain yield and grain color, like the standard carioca were selected, and were intercrossed to generate the population of the following cycle. This process was repeated for four cycles. The best families were evaluated in each cycle by many generations and locations, and the five best lines of each cycle were identified. The 20 lines thus obtained were evaluated in two growing seasons, sown in July and November 2002. The grain yield (kg/ha) and grain type (scale of scores) were evaluated. Genetic progress was confirmed for both traits. The mean annual gain with selection for the grain type was 10.5% and 5.7% for grain yield, with no evidence of variability reduction in the population. These results show that recurrent selection is a good alternative for improving common bean quantitative traits.     

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.     

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.     

Effect of selection for pollen grain size on various traits in common bean

Summary Selection among microgametophytes usually exploits variation in pollen grain germination. Studies of variation in pollen grain size in common bean (Phaseolus vulgaris L.) suggested that selection for size might lead to changes in sporophytic traits. To determine whether microgametophytic selection based on size would affect pollen grain size in subsequent generations or sporophytic traits that were correlated with pollen grain size, pollen grains from three crosses were separated into two size categories by sieving and then used to pollinate cv. Diacol Calima. Selection resulted in changes in pollen grain diameter for pollen from F₁, F₂ and F₃ plants for all crosses. In vitro germination indicated no differences between vigor of large and small grains, but extraction and sieving reduced germinability. F₁ seed from two of the crosses with size-selected pollen varied in weight according to pollen grain size, but in subsequent generations, the effect disappeared. Both size categories of selected pollen resulted in F₂ progeny with reduced numbers of seeds per pod as compared to controls, suggesting that the size selection process may have resulted in indirect selection for traits reducing seed set. The overall results suggested that genes determining pollen grain size in bean have little or no effect on sporophytic traits such as seed size and seed yield.     

Inter-and intraracial hybridization and selection for seed yield in early generations of common bean,Phaseolus vulgaris L.

Summary Five parents from each of four race groups of common bean (Phaseolus vulgaris L.) were hybridized to produce five crosses within each group. Also, five crosses were made for each of the six possible combinations among four groups. Parents, F₁ and F₂, and parents, F₂ and F₃ were evaluated for seed yield in 1990 and 1991, respecitively, at two locations in Colombia.Yield of parents belonging to Middle American races and crosses among them was higher than that of races of Andean origin. Positive correlations were found among the mid-parent value, F₁, F₂ and F₃. Also, the mid-parent value predicted the mean seed yield of all possible lines that could be derived from the F-generation in 42 out of 47 crosses. Four crosses, all between common bean races of Andean and Middle American origin, indicated a possible loss of favorable epistatic parental alleles. On average, mean yield of interracial F₁ hybrids was higher than that of intraracial ones.Positive heteroris (26.4%–123.8%) over the mid-parent in 31 crosses, and F₁s yielding higher (23.7%–91.8%) than the high parent in 20 crosses and yielding higher (22.1%–53.2%) than the highest control among all parents (MAM 13) in 12 crosses, were found. Heritability, estimated by the parent-offspring regression, ranged from 0.42± 0.07 to 0.49±0.04. Expected and realized gains from selection (at 20% selection pressure) ranged from 10.3% to 21.0% over the mean of F₁ hybrids and F₂ and F₃ population bulks.     

Comparison of near-isogenic lines for anthracnoseAre are genes in common bean,Phaseolus vulgaris L.

Summary This study was undertaken to determine whether the Are gene controlling anthracnose resistance had an adverse effect on maturity and yield because backcross-derived cultivars/lines had numerically lower yield and later maturity than the respective recurrent parent. Three pairs of common bean lines near-isogenic for theAre gene were developed fromAre are F₂ plants of the sixth backcross of Seafarer, Fleetwood, and Ex Rico 23. The near-isogenic lines were tested along with the recurrent cultivars and backcross-derived cultivars/lines. The results showed no evidence of any adverse effect of theAre gene on the agronomic characteristics in 11 trails at 5 locations in 3 years.     

Inheritance of temperature sensitivity of the photoperiod response in common bean (Phaseolus vulgaris L.)

Summary Photoperiod response of flowering in common bean (Phaseolus vulgaris L.) is thought to be controlled by the genes Ppd and Hr. However, cultivars also vary in the degree that cooler temperatures reduces their sensitivity to photoperiod. To examine the inheritance of this temperature sensitivity, crosses of cvs. Gordo x de Celaya and Flor de Mayo × Rojo 70 were evaluated at two sites differing in mean temperature and using 12.5-h natural photoperiod or 18-h artificially extended photoperiod. Under 18-h photoperiod at the warmer site, Palmira, no plants of the parents or of the F₂ populations flowered, confirming that the parents were sensitive to photoperiod. Under 12.5-h photoperiod at the cooler site, Popayan, the parents for each cross flowered at similar dates and no segregation for days to flower was observed. However, under 18-h photoperiod, de Celaya and Rojo 70 and the F₁ populations did not flower within 100 days after planting, while the F₂ and F₃ populations showed segregation that was consistent with single gene inheritance, late flowering being dominant. Late flowering at Popayan under 18-h photoperiod indicates a lack of temperature sensitivity, so temperature insensitivity of the photoperiod response was dominant to sensitivity. The name Tip, for temperature insensitivity of photoperiod response, is proposed for this gene, with the recessive form of this gene conditioning earlier flowering at cooler temperatures with long daylengths. It is recognized that the observed segregation patterns could represent the effect of multiple alleles at the Ppd or Hr loci, and studies are proposed to test this possibility with molecular markers and recombinant inbred lines.     

Rooting of bean leaves and use in germplasm evaluation for common bacterial blight resistance

Summary A simple protocol for leaf rooting in beans (Phaseolus vulgaris L.) was developed and used to investigate the reaction of Xanthomonas campestris pv. phaseoli (Xcp), causal pathogen of common bacterial blight disease, in detached versus attached bean leaves. Trifoliate leaves of different sizes (one-third, two-thirds, and fully expanded), either with or without the pulvinus attached to the petioles, were excised from 20 day-old plants of six bean cultivars/lines. Leaf cuttings were cultured in potting medium and then incubated for 5 to 10 days under transparent polyethylene plastic cover in the greenhouse. Roots were readily initiated along the petioles of the leaf cuttings, whether the pulvinus was present or absent. All leaves which were two-thirds expanded and fully expanded developed roots 5 to 7 days after culture. Eighty to 90 percent of the leaves which were one-third expanded formed roots 8 to 10 days after incubation. Laminae of the rooted leaf cuttings were viable and green during the 2 to 3 months period in culture after removing the plastic cover. The common bacterial blight reactions were similar for inoculated attached leaves, detached rooted leaves (inoculated either after or prior to rooting), and moistened detached leaves incubated without rooting. The latter were only usable for evaluation of the Xcp reaction in growth chamber experiments but not under greenhouse conditions. The rooted leaves would be useful for screening bean lines for multiple disease resistance, especially if the pathogens require different environments for disease expression.Abbreviations CBB Common Bacterial Blight - Xcp Xanthomonas campestris pv. phaseoli (Smith) Dye     

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.     

Evaluation of a DNA probe for the quantitative detection of common bacterial blight in common bean and its application in a breeding program

Summary Breeding of Phaseolus vulgaris L. for resistance to common bacterial blight (CBB) can be done with visual evaluations of symptoms to distinguish broad resistance classes, but a more quantitative measure was needed for genetic studies of resistance. A novel method of evaluation was developed by quantifying Xanthomonas campestris pv. phaseoli (XCP) in bean leaf tissue infected with CBB using a ³²P-labeled probe and densitometric analysis of hybridization signals. Quantification of bacterial populations using the probe was highly correlated (r=0.98) with the number of colony forming units (CFU) from plate counts of the same leaf samples. The probe was used to follow XCP population dynamics on susceptible (BAT 41) and resistant (OAC 88-1) bean genotypes. OAC 88-1 supported a maximum XCP population which was approximately tenfold less than BAT 41. The probe was also used to study an F₂/F₃ population segregating for resistance. Narrow sense heritability estimates were less for resistance measured on the basis of bacterial populations (0.18–0.26) than on visual scores of symptoms (0.29–0.38). The anticipated response to selection for CBB resistance would be less based on bacterial numbers than based on symptom expression in this population. In breeding for resistance to CBB, selection based on visual symptoms combined with measurements of XCP populations using a DNA probe can be used to develop bean genotypes that are both resistant to symptom development and bacterial multiplication.Abbreviations CBB common bacterial blight - CFU colony forming units - XCP Xanthomonas campestris pv. phaseoli     

Inter simple sequence repeat (ISSR) markers as a tool for the assessment of both genetic diversity and gene pool origin in common bean (Phaseolus vulgaris L.)

In this study, we report the use of ISSR to assess genetic diversity and to determine the relationships among ten cultivars of common bean developed in Argentina and three materials from France. ISSR markers resolved two major groups corresponding to the Andean and Mesoamerican gene pools of common bean. We compared the results of previous analysis, performed with RAPD markers (Galván et al., 2001), with the results generated by means of ISSR. It appears that ISSR are better tools than RAPDs to identify beans by gene pool of origin though they did not revealed as many differences between individuals as RAPDs. This revised version was published online in July 2006 with corrections to the Cover Date.     

Growth analysis in a reduced leaf mutant of common bean (Phaseolus vulgaris L.)

Summary A comparison between the Royal Red bean variety and a reduced leaf mutant on the basis of growth indices is reported. The mutants has lower leaf area index (LAI) and leaf area duration (LAD) (–43.7%), higher specific leaf weight (SLW) (on the mean +33.3%) and accumulates less dry matter (–7.7%) than the normal genotype. When the net assimilation rate (NAR) is considered, the mutant is superior to the normal (+62.3%). It is concluded that the unit area of the mutant leaves accumulates dry matter more efficiently than that of the corresponding normal variety.     

A non-destructive selection method for faster growth at suboptimal temperature in common bean (Phaseolus vulgaris)

Summary A non-destructive method has been developed to select common bean (Phaseolus vulgaris L.) plants whose growth is less effected at a suboptimal temperature. Shoot weight was determined at a suboptimal (14°C) and optimal temperature (20°C), 38 days after sowing and accessions identified with a significantly lower than average weight reduction at 14°C compared to their weight at 20°C. Weight of primary leaves and of the shoot was correlated with seed weight at both temperatures, but no correlation was found between shoot weight reduction at 14°C and seed weight.     

Combining ability for nodulation in common bean (Phaseolus vulgaris L.) genotypes from Andean and Middle American gene pools

Ten F₁'s obtained from crosses among five common bean genotypes of Andean (WAF 15, Mineiro Precoce and Batatinha) and Middle American (BAT 304 and Ouro) gene pools were assessed for their combining abilities for root nodulation with Rhizobium tropici strain CIAT 899. The plants were grown under controlled conditions and evaluated for number of nodules per plant (NN), nodule dry weight (NDW), mean nodule weight (MNW) and plant fresh weight (PFW). The subdivision of the treatment effects on the general (GCA) and specific combining effects (SCA) were performed according to Griffing's diallel analysis method 2, model 1. The analyses of variance and estimates of quadratic components showed that non-additive gene effects were more important in the expression of NN and PFW, whereas additive gene effect was predominant for NDW and MNW. A close association was observed between high number of nodules and GCA. Generally, crosses involving parents of different gene pools yielded hybrids with high positive estimates of SCA for all assessed traits. The genotypes of Andean origin WAF 15 and Mineiro Precoce are the most promising parents for breeding programs to increase NN and NDW in common beans. This revised version was published online in July 2006 with corrections to the Cover Date.     

Natural selection for grain yield in dry bean populations bred by the bulk method

Three dry bean harvests are possible in some regions of Brazil that differ significantly in temperature, rainfall and day length conditions. Thus in breeding programs, generation advance using the bulk method can be carried out in these three periods. It is questionable whether under conditions such as these the action of natural selection would contribute to an increased frequency of individuals considered superior by breeders. The present study assessed six segregating populations, for up to 17 successive generations to check the effects of natural selection during inbreeding. Six populations were generated from a partial diallel: one group consisting of two early maturity cultivars, ESAL 686 and Manteigão Fosco was crossed to a second group of three cultivars with medium maturity, Carioca MG, Milionario and Ouro. The six populations plus the five parents were assessed in experiments using a randomized complete block design, in three locations: Lavras, where in breeding started from the F₂ generation; Lambari and Patos de Minas, from the F₃ generation. A linear regression equation was fitted to the parents and hybrids mean data in each location, considering grain yield as the dependent variable (y) and generations (F₂ = 1, F₃ = 2, etc.) as the independent variable. Genetic progress was estimated from the differences between the linear regression coefficients of the hybrids (b_i)and the mean of the b coefficient of their respective parents (b_j). The (b_i-b_j) were positive in all cases, showing that natural selection acted in all the segregating populations and contributed to an average increase in grain yield of 2.5% per generation over the mean of the parents.