Inheritance of seed coat colour in broad bean (Vicia faba L.)

Summary The inheritance of the seed coat colours violet, spotted, brown, green, red, black and beige was investigated in a 10×10 diallel cross between broad bean (Vicia faba L. major) lines. Spotted seed colour was dominant over any uniform seed coat colouring. Brown was dominant over black, green and normal (beige colour). Black and red seed parents behaved as recessive in all F₁ progenies. A 3 (coloured): 1 (normal) segregation ratio was observed in the F₂ of crosses of violet, brown, black, red and spotted seed coat parents to nornal seed coloured parents. Green x beige gave a segregation ratio of 9:7 in F₂. When two parents with different seed coat colour were involved in a cross, the F₂ showed a typical digenic segregation ratio thus demonstrating two unlinked and sometimes epistatic loci.Segregation of a multiallelic series at two loci explains all segregation ratios observed for seed coat colour in broad bean.     

Genetic linkage map of Chinese native variety faba bean (Vicia faba L.) based on simple sequence repeat markers

Simple sequence repeat (SSR) marker is a powerful tool for construction of genetic linkage map which can be applied for quantitative trait loci (QTL) and marker‐assisted selection (MAS). In this study, a genetic map of faba bean was constructed with SSR markers using a 129 F₂ individuals population derived from the cross of Chinese native variety 91825 (large seed) and K1563 (small seed). By screening 11 551 SSR primers between two parents, 149 primer pairs were detected polymorphic and used for F₂ population analysis. This SSR‐based genetic linkage map consisted of 15 linkage groups with 128 SSR. The map encompassed 1587 cM with an average genetic distance of 12.4 cM. The genetic map generated in this study will be beneficial for genetic studies of faba bean for identification of marker‐locus‐trait associations as well as comparative mapping among faba bean, pea and grasspea.     

Inheritance of seedling colour in faba bean (Vicia faba L.)

Summary The inheritance of purple seedling colour was studied, in relation to the genetic control of flower colour. It was found that purple seedling colour is likely to be controlled by a single gene and that the trait is dominant over green seedling colour. White flowering prohibited the expression of the purple seedling colour, and is therefore thought to be epistatic.This character can be used to estimate rate of outcrossing in breeding programmes, as well as contribute to our knowledge of the biosynthesis of plant pigments and secondary metabolites such as tannins.     

Rate and inheritance of cross-fertilization in faba bean (Vicia faba L.)

Summary To assess the feasibility for synthetic varieties of Vicia faba L., 108 faba bean genotypes were screened for cross-fertilization rate in five field experiments carried out in 1989 and 1990 at Wageningen.Significant genotypic differences were observed for outcrossing rate, which ranged from 1 to 55%. These percentages, however, are thought to be not high enough for a commercially feasible production of synthetic varieties.The inheritance of cross-fertilization rate, determined by testing the F₄ generation of a cross between two inbreds, both with high outcrossing rates, appeared to be of a polygenic nature, as was indicated by the wide variation in, and the continous distribution of, the cross-fertilization rates. Among the F₄ lines tested some had a significantly higher cross-fertilization rate than the parents. However, this difference was not found when these lines were retested, indicating that possibilities for successful selection for higher outcrossing rates might be limited.     

Single gene mutation for green cotyledons as a marker for the embryonic genotype in faba bean, Vicia faba

A new spontaneous mutation is described in faba bean, Vicia faba L., characterized by a marked green colour in the entire cotyledon tissues of the mature seeds, while the wild phenotype is yellow. This seed character reflects the embryonic genotype and is determined by a single recessive allele named i1-1. It is distinct from the y gene, which codes for green testa. As is the case for gene y, the gene i1-1 displays no epistatic effects with the zero-tannin genes zt1 and zt2 which influence seed and flower colour in faba bean. In this allogamous species, such a mutation is a useful tool in cross-fertilization studies.     

Heterosis and hybrid performance in topless faba beans (Vicia faba L.)

Parental lines, F₁ and F₂ hybrids from a 7×7-diallel of topless faba beans and eight standard cultivars were grown in single-rows for two years at four sowing dates on a highly fertile loam soil near Göttingen. In grain yield, F₁ hybrids showed 21 to 54% advantage over the higher yielding parent. Compared with the midparental value an average heterosis of 50% was observed for grain yield. Corresponding heterosis mean values ranged from 3 to 37% for yield components and up to 11% for phenological traits. The grain yield of the highest yielding topless F₁ hybrid was equal to that of Alfred, the highest yielding indeterminate standard cultivar. High grain yield in the topless hybrids was closely correlated with high biomass yield, late ripening and tall plants generally having a high number of podded nodes and pods on the main stem. Due to their agronomic advantages, i.e., lower plant length, reduced lodging sensitivity and earlier ripening, efforts at breeding topless hybrids seemed to be worth-while in faba beans. However, their yielding ability must be improved further before they can compete with normal, semi-dwarf and stiff-strawed cultivars.     

Development of EST-SSR markers and construction of a linkage map in faba bean (Vicia faba)

To develop a high density linkage map in faba bean, a total of 1,363 FBES (Faba bean expressed sequence tag [EST]-derived simple sequence repeat [SSR]) markers were designed based on 5,090 non-redundant ESTs developed in this study. A total of 109 plants of a ‘Nubaria 2’ × ‘Misr 3’ F₂ mapping population were used for map construction. Because the parents were not pure homozygous lines, the 109 F₂ plants were divided into three subpopulations according to the original F₁ plants. Linkage groups (LGs) generated in each subpopulation were integrated by commonly mapped markers. The integrated ‘Nubaria 2’ × ‘Misr 3’ map consisted of six LGs, representing a total length of 684.7 cM, with 552 loci. Of the mapped loci, 47% were generated from multi-loci diagnostic (MLD) markers. Alignment of homologous sequence pairs along each linkage group revealed obvious syntenic relationships between LGs in faba bean and the genomes of two model legumes, Lotus japonicus and Medicago truncatula. In a polymorphic analysis with ten Egyptian faba bean varieties, 78.9% (384/487) of the FBES markers showed polymorphisms. Along with the EST-SSR markers, the dense map developed in this study is expected to accelerate marker assisted breeding in faba bean.     

The identification of faba bean (Vicia faba L.) trisomics by translocation tester set

Summary Five subtelocentric primary trisomics of Vicia faba L. were isolated from crosses between asynaptic mutants and normal plants or translocation tester stocks. The extra chromosome of each trisomic was identified cytologically by critical multivalent association at metaphase I of meiosis.     

Pollination,fertilization and seed development in winter stocks of faba beans (Vicia faba L.)

Summary Incidences of pollination and fertilization were determined in five stocks of winter bean (two populations, two inbred lines, one cultivar) in two years. In the second year, irrigation was included as an experimental treatment and the development of seeds was monitored. Large differences between years were found in the measures of pollination and fertilization, and irrigation was associated with a small but significant increase in the proportion of ovules which were fertilized within fertilized flowers, particularly in the ovules furthest from the stigma. This increase was clearly reflected in the corresponding indices of seed maturation in the irrigated plants. Irrigation was not associated with any change in the proportion of ovules which aborted. Small but statistically significant differences in all of the measures of pollination and fertilization were found among the stocks. It is suggested that the variation in the incidences of pollination and fertilization was largely associated with variation in the synchrony of anthesis within racemes. The proportion of ovules within pods which matured as seeds and the proportion which aborted both varied significantly among the five stocks. Abortion appeared to act independently of fertilization in affecting the index of seed maturation, so both mechanisms should be considered when breeding for improved filling of pods.     

Effectiveness of early generation selection under two plant densities in faba bean (Vicia faba L.)

The effectiveness of early generation selection under low and high plant density was evaluated in faba bean (Vicia faba L.). For this, the faba bean cv. 'Polycarpe' was pollinated by eight higher yielding cultivars of different origin. The F₁'s and F₂'s together with 'Polycarpe' and the pollinator with the highest yielding ability (cv. 'Effe') were evaluated under high plant density with adjacent controls and under low plant density in a honeycomb design. It was found that the eight hybrids had a different ranking at low than at high plant density for both generations (F₁and F₂). Hybrid Pol.x A-58 had the best combined yield performance in the F₁and F₂under high and hybrid Pol.x A-37 under low plant density. Application of individual plant selection in the F₂and F₃generation of the two crosses (Pol.x A-58 and Pol.x A-37), under both low plant density according to UNR-O honeycomb design and high plant density using a moving average, resulted in the selection of four groups with 20 selected F₃plants each. Within each group, equal numbers of seeds from each plant were mixed to establish four mixtures. These mixtures together with two controls ('Polycarpe' and 'Effe') were evaluated in a randomized complete block design (22 plants/m²) in one location for two years. It was found that the material selected continuously under low plant density had a significantly higher yield than the material selected under high plant density. In addition, the material selected from Pol.x A-37 (H₁R₂) under high plant density had a lower mean yield than that selected continuously under low plant density (H₁H₂). Their average yield over years was significantly higher than the yield of the material selected from Pol.x A-58 under both plant densities. Furthermore, the material selected from Pol.x A-58 under low plant density, had a significantly higher mean yield over years than its sibs selected under high plant density. It was concluded that honeycomb selection at low plant density can be used effectively for early generation selection in faba bean.     

Intraspecific Variation in Mean Endosperm Cell Cycle Time in Vicia faba (L.) and Interspecific Hybridization with Vicia narbonensis (L.)

The rate of early embryo-sac development was studied in seven faba bean (Vicia faba L.) cultivars grown in a controlled environment. The embryo-sac stage was determined from the number of endosperm nuclei per embryo-sac and the number of embryo cells during the first 12 days after pollination. Differences in early embryo-sac development were observed between the genotypes studied. In addition, five different V. faba×V. narbonensis crosses and reciprocals were made using genotypes with small differences in mean endosperm cell cycle time and genotypes with large differences. The percentage of success (pod set) in V. faba×V. narbonensis crosses ranged between 9 % and 59 % while in the reciprocals it ranged from 12 % up to 30 %. However, cytological studies showed that the high percentage of success (pod set) observed in the cross A-107 × A-202 was not associated with a higher percentage of interspecific hybrid embryos. The results indicate that genotypes of the two species with smaller differences in mean endosperm cell cycle time result neither in a higher percentage pod set nor in bigger hybrid embryos.     

Pattern analysis and genotypic x environment interactions in faba bean (Vicia faba L.) populations

Summary Twenty-nine populations of faba bean (Vicia faba L. subsp. major, equina and minor) of mediterranean origin were evaluated for the genetic variability of 12 biomorphological characters at four locations in Apulia and Basîli cata regions (Italy) in 1983–84.A pattern analysis appoach, based on classification and ordination procedures is presented for to characterize differences among population in mean performance and response across environments. Informations about G × E interaction were obtained.Research work supported by CNR, Italy. Special grant I.P.R.A. Sub-project 1. Paper N. 1099.     

Heterosis and inheritance of drought tolerance in faba bean, Vicia faba L.

Twelve parental lines of faba beans and 27 F₁ hybrids derived from them were field-tested under a dry and a well-watered treatment, at two German locations (Göttingen, Hohenheim) in two years (1995, 1996). Drought was artificially induced using rain shelters from anthesis until maturity. Data were collected on yield under dry (Yd) and well-watered (Yw) conditions. Drought tolerance was assessed as Yd/Yw. The mid-parent heterosis was significant for Yd and Yw, but not for Yd/Yw. Relative heterosis for Yd (52.0%) was greater than for Yw (39.3%). Parental heritability was greater for Yw (0.86) than for Yd (0.61). However, it was similar for both traits in F₁ hybrids (0.65 and 0.66). Principal component analysis showed that F₁ hybrids exhibited the same pattern of adaptation as their parents. The heritability of drought tolerance was 0.48 in F₁ hybrids and 0.70 in parents. Attention should be paid to its negative correlation with yield. Several hybrids combined high tolerance, a favourable pattern of adaptation and appropriate performance.     

Density enhancement of a faba bean genetic linkage map (Vicia faba) based on simple sequence repeats markers

Genetic mapping for faba bean lags far behind other major crops. Density enhancement of the faba bean genetic linkage map was carried out by screening 5,325 genomic SSR primers and 2033 expressed sequence tag (EST)‐SSR primers on the parental cultivars '91825' and 'K1563'. Two hundred and fifteen genomic SSR and 133 EST‐SSR primer pairs that detected polymorphisms in the parents were used to screen 129 F₂ individuals. This study added 337 more SSR markers and extended the previous linkage map by 2928.45 cM to a total of 4516.75 cM. The number of SSR markers in the linkage groups varied from 12 to 136 while the length of each linkage group ranged from 129.35 to 1180.21 cM. The average distance between adjacent loci in the enhanced genetic linkage map was 9.71 cM, which is 2.79 cM shorter than the first linkage map of faba bean. The density‐enhanced genetic map of faba bean will be useful for marker‐assisted selection and breeding in this important legume crop.     

Study on the Inheritance and Relationship of Some Morphological and Fertility Characters in Faba Bean (Vicia faba L.)

Inheritance of several morphological and fertility characters was studied in F2 generation of two sets of diallel cross, using Griffing's analysis of variance. A paramount importance of general combining ability (GCA) over specific combining ability was measured for most studied characteristics. Correlations among GCA values for seed yield, morphological and fertility characters do not give a clear definition of the plant ideotype. “Terminal inflorescence” character appeared to reduce the seed per ovule ratio. Higher additive component was found for the ovule per pod than for seed per pod characters.     

A xenia effect on number and volume of cotyledon cells and on seed weight in faba bean (Vicia faba L.)

The female line 1709, carrying the recessivecharacter green cotyledon, marker of the embryogenotype, was cross-pollinated by bees in the presenceof either male line HG115 or male line Dial63. In thecomparison of seeds which resulted either fromintercrossing or from selfing, a xenia effect wasobserved on volume and number of cotyledon cells andon individual seed weight which was dependent on themale line. The increase of hybrid seed weight inducedby the small-seeded male line Dial63 was larger thanthe one induced by the large-seeded male line HG115.The xenia effect on seed weight was correlated withthe level of heterosis in the F1 plant's growth.     

Untersuchungen zur Leitbündelstruktur der Ackerbohne (Vicia faba L.)

Investigations on the vessel structures of broad beans ( Vicia faba L.)
The number of vessels was observed in the fruit bearing axes of some varieties from Vicia faba Minor, Equina and Major during flowering. These investigations were also made on some F₂- progenies of crosses from minor × equina and Paucijuga × Eufaba and on some topflower plants.
The varieties of Minor and Equina had about 12 vessels on the base of the axis, these diminished to the top of the axis 3 or 4 vessels just before the last flower. The varieties of Major had 12 vessels on the base of the axis too, but these diminished faster because of the smaller number of flowers on the axis. In Paucijuga the number of vessels on the base of the axis was only 3 to 4. Plants of Paucijuga had only 2 or 3 flowers per axis.
Progenies from crosses between Minor × Equina had more vessels on the base of the axis, probably due to a heterotical inheritance. In progenies from crosses between Paucijuga × Eufaba an intermediate inheritance was found in the number of flowers per axis and in the number of vessels.
Due to the investigations on a comprehensive material it is probable that the described vessel structure exists in most broad beans. This structure is complex arranged and modifyable. With this organization it is quite unprobable that the vessel structure will be a yield reducing factor.     

Detection and characterization of two new CMS systems in faba bean (Vicia faba) *

Wide crosses were made to identify new cytoplasmic male sterility (CMS) systems in faba beans, based on the interaction of cytoplasm with restorer and maintainer alleles. A total of 330 F₁ hybrids were produced in both reciprocal forms. Male sterile segregates were observed in one reciprocal version in the F₂ generation of six crosses. Two of these crosses with female parents originating from Afghanistan and Egypt expressed stable male sterility in subsequent backcross generations. Based on the female parents of the two crosses, these two CMS systems were designated CMS 199 and CMS 297. CMS 199 was more stable than CMS 297 during backcross generations and across different environments. Maintainer and restorer lines for both CMS systems were identified. Lower expression of male sterility occurred in CMS 297 in the greenhouse during the winter generations than in isolation cages during the summer generations, which may be utilized to maintain male sterile lines by selfing. Regarding practical applications, the CMS 199 shows great promise for hybrid breeding in faba beans.     

New Isozyme Markers in Vicia faba: Inheritance and Linkage

Results on the inheritance of 6 enzyme systems: LDH, PGM, FDH, SKD, SOD, AAT from seeds of Vicia faba and the linkage relationships among these isozyme loci are presented. The allozymes at each one of these loci behaved in a codominant manner and segregated in the expected Mendelian ratio. Linkage tests between these loci showed that they segregate independently.     

Differential Drought Responses of Faba Bean (Vicia faba L.) Inbred Lines

Drought responses of 19 inbred faba bean lines of different origin were studied in the field under rain shelters with and without irrigation. Inbred lines differed significantly in response to drought (P < 0.01): those with a lower drought sensitivity index (SI) (more resistant) originated from the drought-prone regions characterized by smaller plant size (r = 0.93), and more pods and seeds per plant (r ≥ 0.90) regardless of seed size, while lines with higher SI (more sensitive) were those which mainly exhibited higher yield under favourable conditions accompanied by a greater biomass. In a greenhouse experiment under mild drought (−0.15 MPa soil water potential), comparisons between relatively drought-sensitive (Adriewaalse) and drought-resistant (L7) inbred lines showed that Adriewaalse used 38 % more water than L7 and also produced 40 % more biomass. There was a stress-induced decrease in osmotic potential (ψ_s) in both lines (by 0.72 and 0.50 MPa for Adriewaalse and L7, respectively) accompanied by decreased turgor in Adriewaalse and increased turgor in L7. The difference in drought-induced solute accumulation between lines was diminished when solute accumulation due to water loss and growth inhibition was considered, which indicates that solute accumulation was the result of a concentration effect. Similarly, lower SI in the field was not the result of osmotic adjustment, as the relationship between SI and drought-induced decrease in ψ_s was negative. The water use efficiency of both inbred lines increased markedly with increasing water deficit, though there was no difference between the lines. It was concluded that differences in drought resistance between these inbred faba bean lines were manifested through plant size-induced water demand (avoidance) but were not associated with osmotic adjustment (tolerance).