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.     

Genetics of resistance to bean golden mosaic virus in Phaseolus vulgaris L.

Summary The genetics of resistance to bean golden mosaic virus (BGMV) was studied in an 8×8 complete diallel cross of common bean, Phaseolus vulgaris L. The 28F₁ hybrids, their reciprocals, and eight parents were artificially inoculated with BGMV under glasshouse conditions. Data were recorded and analyzed for foliar yellowing, plant dwarfing, flower abortion, and pod formation, using a randomized complete block design with two replications.General combining ability (GCA) mean squares were highly significant (P<0.01) and larger than values for specific combining ability for all traits. Reciprocal and maternal effects were nonsignificant. Cultivars Royal Red and Alubia Cerrillos possessed significant negative and positive GCA for all traits, respectively. Porrillo Sintetico also had negative GCA for all traits except plant dwarfing. None of the GCA effects were significant for Great Northern 31 and PVA 1111. Positive associations existed among all traits studied.     

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.     

Inheritance of the mosaic and necroses reactions induced by bean severe mosaic comoviruses in Phaseolus vulgaris L.

The inheritance of the localized necrosis, apical necrosis, and mosaic reactions induced by bean severe mosaic comoviruses in common bean (Phaseolus vulgaris L.), was studied in crosses of Great Northern 123 × Pitouco, Great Northern 123 × Iguaçu and Pitouco X Iguaçu. Great Northern 123 reacts with mild mosaic, Iguaçu with localized necrosis, and Pitouco with apical necrosis to bean severe mosaic comoviruses. The analysis of the F₁ and F₂ generations indicated that localized necrosis was dominant over mild mosaic, and apical necrosis was dominant over the localized necrosis and the mild mosaic. An independently inherited single dominant gene controlled the expression of localized and apical necrosis in cultivars Iguaçu and Pitouco, respectively. The dominant gene for apical necrosis (Anv) found in Pitouco, was epistatic over the dominant gene (Lnv) conditioning localized necrosis in Iguaçu, as suggested by the F₂ of Pitouco × Iguaçu, that yielded a ratio of 12 apical necrosis: 3 localized necrosis:1 mild mosaic. The genotypes of Pitouco, Iguaçu and Great Northern 123 are: Anv Anv lnv lnv, anv anv Lnv Lnv, and anv anv lnv lnv, respectively.     

A rapid screening technique to combine resistance to halo blight and bean common mosaic virus in Phaseolus vulgaris L.

Summary Seven bean lines (Phaseolus vulgaris L.) with differential resistance or susceptibility to race 2 of halo blight (Pseudomonas syringae pv. phaseolicola) and a necrosis-inducing isolate of bean common mosaic virus were inoculated with one or both pathogens in combination, to determine the feasibility of dual screening to identify resistance to both pathogens simultaneously. Dual screening yielded the same results as separate screenings. Neither pathogen affected the disease expression of the other. Simultaneously screening for resistance to both pathogens will shorten the recurrent screening-selection cycle of hybridization programs, and accelerate development of resistant cultivars.Abbreviations BCMV Bean Common Mosaic Virus - cvs Cultivars - HB Halo blight - Inoc. Pt. Inoculation point - NLL Necrotic local lesion - React Reaction - SVN Spreading veinal necrosis, System chloro-Systemic chlorosis - VN Vascular necrosis     

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.     

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.     

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.     

Yield stability in common bean (Phaseolus vulgaris L.) genotypes

An experiment was undertaken to determine the stability of seed yield in 21 common bean genotypes representing three growth habits. Seven genotypes in each growth habit (determinate bush, indeterminate bush and indeterminate prostrate) were evaluated in replicated trials at three locations for three years under rain fed conditions in Ethiopia. A combined analysis of variance, stability statistics and rank correlations among stability statistics and yield-stability statistic were determined. The genotypes differed significantly for seed yield and genotype × environment (year by location) interaction (GE). The different stability statistics namely Type1, Type 2 and Type 3 measured the different aspects of stability. This was substantiated by rank correlation coefficient. There were strong rank correlations among S_i ²d, W_i ²,σ_i ² and S_i ², where as there was weak correlation between b_iand R_i ², S_i ²d, W_i ², σ_i ² and S_i ². R² was significantly and negatively correlated with W_i ², σ_i ² and S_i ². σ_i ² is significantly correlated with W_i ².Yield is significantly correlated with b_i and R_i ².None of the statistics per se was useful for selecting high yielding and stable genotypes except the YS(yield-stability statistic). Most of the high yielding genotypes were relatively stable. Of the 21 genotypes, only 11genotypes were selected for their high yielding and stable performance. Genotypes with growth habit III and I (in determinate prostrate and determinate bush) were generally more stable than in determinate bush. This revised version was published online in July 2006 with corrections to the Cover Date.     

Male sterility in beans (Phaseolus vulgaris L.)

Summary Male sterile bean plants were discovered at Embu Agricultural Research Station, Kenya. The character is controlled by one recessive gene.The Grain Legume Project is a joint project of the Kenya Ministry of Agriculture and The Netherlands Ministry of Foreign Affairs.     

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.     

Alternative methods of interpreting measurements of resistance to the leafhopper Empoasca kraemeri Ross and Moore in the common bean,Phaseolus vulgaris L.

Summary Appropriate definitions of, and methods of measuring, resistance are sought using Phaseolus vulgaris, a major source of protein in Latin America, and its most important pest, Empoasca kraemeri. The concepts of general vigour and narrow-sense resistance are defined: these two components make up broad-sense resistance. The validity of these definitions, which depend on the inclusion of tolerance as a resistance mechanism, is discussed. In order to distinguish the components, 38 genotypes of P. vulgaris were grown with and without insecticidal proctection, in three replications in a split-plot design, with three planting dates. Seed yield per plant, and a visual score of damage symptoms or vigour, were measured. Three methods of estimating narrow sense resistance are considered: 1) the genotype × insecticide interaction effect in a split-plots analysis, 2) the yield of each genotype unprotected (Y_u) divided by its yield protected (Y_p). and 3) the deviation of each genotype from a regression of Y_u on Y_p. The third method overcomes some objections to the first two, and can be applied to damage and vigour scores as well as to yields. The 38 genotypes differed in narrow-sense resistance according to all three methods, and it appears that in practice Methods 2 and 3 were both valid. They also differed in general vigour. Breeders working with many pests and crops have found that low damage and high yield in the presence of the pest are effective selection criteria. The work reported here supports this view.     

Variation of seed storage proteins in landraces of common bean (Phaseolus vulgaris L.) from Basilicata,Southern Italy

Summary Twenty landraces of common bean (Phaseolus vulgaris L.) from Sarconi and Rotonda two locations of Basilicata, a Southern Italy region, were screened for variation in seed storage proteins (phaseolin and phytohemagglutinin) by polyacrylamide gel electrophoresis (SDS/PAGE and IEF-SDS/PAGE). No variation of the main seed protein fractions was observed within each landrace. Phaseolin patterns type C and T were exhibited from the landraces; the type C resulted predominant with a frequency of 70%. Only the C type was observed for the landraces of Rotonda, T and C for those from Sarconi. Two variants were observed for the phytohemagglutinin by SDS-PAGE, one was common to eighteen landraces. However, these variants submitted to IEF-SDS/PAGE resulted similar to the type T_G2described by Brown. The importance to safeguard these landraces is stressed by the observation that the more spread commercial cultivars of common bean growing in Southern-Italy had a T phaseolin pattern. A reduction of C type diffusion could produce the loss a typical trait of common bean in Mediterranean regions.     

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.     

Effect of plant density on selection for seed yield in two population types of Phaseolus vulgaris L.

Summary Pedigree selection for seed yield, using early generation yield tests, was practiced from the F₂ to F₇ in two populations of common bean (Phaseolus vulgaris L.) in three plant densities: 66, 133, and 266 thousand plants ha^-1 at CIAT-Palmira, Colombia. The six highest yielding lines selected from each plant-density, along with 13 parents, were evaluated in a 7×7 lattice design with three replications at the three densities utilized for selection, and at 399,000 plants ha^-1 in 1988 and 1989.Based on the mean performance of selected lines and the mean of the parents, selection for seed yield was effective in all densities in both populations. However, none of the lines selected from the population within the race Mesoamerica (TC 4673) significantly outyielded their best parent under any plant density. The highest yielding lines selected from the interracial population (TR 4635) outyielded their best parent irrespective of the plant-density used for selection. The highest yielding line originated from the highest density used for selection. Low density was neither good for selection nor for evaluation and identification of high-yielding cultivars of common bean. There was no significant difference between the mean yield of lines selected at the intermediate and high population densities. The effects of plant density, year, and their interactions were significant for seed yield.     

Variation of the storage protein phaseolin in common bean (Phaseolus vulgaris L.) from the Mediterranean area

Summary Variation of phaseolin, the major storage protein in Phaseolus vulgaris L., was analysed in seeds of a germplasm collection from the Mediterranean area. Results showed a number of subvariants differing from the reference patterns for the presence, absence or faintness of single polypeptides. Maximum concentration of variation was found in the Cyprus material. Sources and causes of variation are discussed.     

Genetic control of low temperature tolerance in germination of the common bean (Phaseolus vulgaris L.)

Summary Cultivars tolerant to low temperature during the germination and emergence stages and carriers of the grain quality standards demanded by consumers are needed to increase the success of irrigated common bean in Southern Minas Gerais State. To study the genetic mechanisms controlling these traits and assess the possibility of generating the desirable materials, a diallel cross involving ten cultivars including introductions and pure lines from the breeding program of ESAL was carried out. Speed of germination of the F₂ generation from the crosses was assessed under laboratory conditions at 12°C. The diallel analysis was carried out using Griffing's method IV (1956) and predictions of the cross potential were made by the methodology developed by Jinks & Pooni (1986). The results indicated that the parents differed in germination speed at low temperatures with Small White, A-488 and Rio Vermelho being the most tolerant and Carioca, ESAL 591 the most sensitive. No effect of the reciprocal crosses was observed either for germination percentage or germination speed. The parents A-488, Small White and Rio Vermelho showed the greatest general combining ability. Additive genetic effects predominated for both traits. Our results suggest that cold tolerance can be bred successfully into commercially acceptable cultivars.     

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.     

Traits associated with dry edible bean (Phaseolus vulgaris L.) productivity under diverse soil moisture environments

Two experiments were conducted in the Rift Valley, Ethiopia (8°N and 39°E) to determine associations between eight plant traits and seed yield, and to obtain estimates of narrow sense heritability for the traits. Experiment I evaluated seven dry edible bean cultivars/lines at two locations to simulate different soil moisture stress, including, Debre Zeit(non-stress) and Dera (moderate-stress). Experiment II evaluated 25 cultivars/lines in three environments including, Melkassa early planted (non-stress), Melkassa late planted (high-stress), and Dera (moderate-stress). A randomized-complete-block design with three replicates was used in both experiments. Plant traits evaluated were seed yield, pods plant^-1, seeds pod^-1, 100 seed weight, root dry weight, hypocotyl diameter, plant biomass, plant height and days to flowering. Plant traits that were significantly associated with seed yield were included in a stepwise-regression model to determine which trait or combination of traits provided the best model to estimate seed yield in each environment. An analysis of variance was conducted to test main effects and interactions between plant traits and environments. Significant variation among lines occurred for seed yield and all plant traits in both experiments. Strong positive correlations were observed between plant biomass and seed yield in all environments. Seed yield and pods plant^-1 were also highly associated in four of the five environments. Stepwise regression models indicated that the combination of pods plant^-1 and plant biomass consistently contributed to seed yield prediction, while other traits did not. Because both plant biomass and pods plant^-1 had moderate to high narrow sense heritability estimates and low GE interactions, they should be useful as indirect selection criteria to improve and stabilize seed yield in a breeding program. This revised version was published online in July 2006 with corrections to the Cover Date.     

The inheritance of resistance to Empoasca kraemeri Ross & Moore in the common bean,Phaseolus vulgaris L.

Summary Genotypes of Phaseolus vulgaris differ in their level of resistance to Empoasca kraemeri, but the speed with which these differences can be accumulated into a resistant variety will depend on whether they can be detected in single F₂ plants, and on whether the genetic control of resistance is additive. Thirty-eight homozygous genotypes were therefore grown in small plots with and without insecticidal protection, and also in simulated F₂ populations containing one plant of each genotype. In addition, the F₂ and F₃ generations of crosses were studied. Both generations of a 13 parent diallel were grown with and without insecticidal protection. Six crosses, involving some of the parents in the diallel and two additional genotypes, were studied in more detail. Protected and unprotected F₂ plants from these crosses were harvested individually, and the resulting F₃ families were grown with and without protection. In all the expriments, the level of damage to the unprotected plants and the vigour of the protected plants was assessed visually by means of a numerical score, and the seed yield was measured.The performance of a single homozygous plant was a reasonable predictor of the performance of the same genotype in a small plot, but in the six crosses studies in detail, the correlations between F₂ and F₃ performance were low. In the analysis of the diallel cross, the significant differences among crosses were due to specific combining ability and not general combining ability. These results indicate that the genetic control of resistance is largely non-additive, and that F₂ selection will probably be ineffective. However, F₃ selection should be effective, and transgressive segregants for resistance should be obtainable.In some of the six crosses, the F₃ families differed both in general vigour and in narrow-sense resistance to E. kraemeri, as defined by Galwey & Evans (1982). The visual assessment of damage symptoms in each plot was related to the subsequent seed yield of the plot, but was more heritable than yield. The relationship between these two variables was investigated by the genetic correlation coefficient and by an alternative method, and was found to be partly genetically determined, but partly due to the influence of environmental factors on both variables. This indicates that screening for E. kraemeri resistance should be done by means of a visual damage score in a carefully chosen, uniform environment.