Combining ability analysis of resistance to Helminthosporium leaf blight in spring wheat

Wheat breeders in South Asia are attempting to develop wheat (Triticum aestivum L.) cultivars resistant to Helminthosporium leaf blight (HLB), which occurs mainly as a complex of spot blotch caused by Cochliobolus sativus (Ito & Kuribayashi) Drechs. ex Dastur, and tan spot caused by Pyrenophora tritici-repentis (Died.) Drechs. Information on the combining ability for HLB resistance in wheat cultivars of South Asia is not available. This study was undertaken to examine the resistance to HLB in nine genetically diverse wheat parents, and to evaluate their general combining ability (GCA) and specific combining ability (SCA) effects toward determining the genetic basis of disease resistance. Nine parents were crossed in a half-diallel mating design to produce 36 populations. The F₁ and F₂ progenies, and the parents were evaluated in replicated field tests at Rampur, Nepal. Multiple disease scores were recorded, and area under the disease progress curve (AUDPC) was calculated to measure disease severity over time. The combining ability analysis was performed using Griffing's Method 2, Model 1. The parents chosen showed wide variation for resistance to HLB. They and the F₁ and F₂ progenies differed significantly for AUDPC. GCA and SCA effects were significant in both generations suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. Wheat genotypes 'SW89-5422', 'G 162', 'NL 781'and 'Chirya 7' had significantly negative GCA effects for AUDPC in both F₁ and F₂ generations, suggesting their prime suitability for use in wheat breeding programs to improve resistance to HLB. The estimate of narrow-sense heritability was 0.77 in both generations suggesting that selection for HLB resistance should be effective in these crosses. The results indicate a predominance of additive gene action in the inheritance of HLB resistance in spring wheat.     

Diallel analysis of resistance to head blight caused by Fusarium culmorum in winter wheat

Summary Ten homozygous winter wheat genotypes representing different levels of resistance to Fusarium head blight were crossed in all possible combinations excluding reciprocals. Parents, F₁ and F₂ were inoculated with one pathogenic strain of Fusarium culmorum. Data for head blight, observed 21 days after first inoculation (OBS-2), and for the area under the disease progress curve, based on observations 14, 21 and 28 days after first inoculation (AUDPC), were analyzed. The contrast between parents and F₁ crosses indicated dommance effects of the resistance genes. Diallel analysis according to Griffing's Method 4, Model 1 showed significant general combining ability (GCA) effects for both F₁ and F₂; specific combining ability effects were not significant. With the exception of one genotype for which general performance for Fusarium resistance was not in agreement with its GCA, the resistance to F. culmorum was uniformly transmitted to all offspring, and the parents can be described in terms of GCA. It is suggested that in the progenies with one of the awned lines as parent, one resistance gene was linked with the gene coding for presence of awns, located on chromosome 4B. A single observation date, taken at the right time, was as effective in assessing resistance as the AUDPC.     

The breeding values of potato parents for field resistance to late blight measured by whole seedlings

Summary Parents used in a potato breeding programme were intercrossed in a multiple mating scheme. The resulting progenies were assessed for resistance to late blight, using as criteria the size and frequencies of lesions on leaves, on petioles and on stems, and an overall score. Analysis of variance showed that all the statistically significant genetic variation was attributable to general combining ability (GCA) differences. The seven variates were highly correlated.     

Diallel analysis of barley for resistance to leaf stripe and impact of the disease on genetic variability for yield components

Barley breeders in Syria attempting to develop barley (Hordeum vulgare L.) cultivars resistant to barley leaf stripe (BLS) disease caused by Pyrenophora graminea Ito & Kuribayashi [anamorph Drechslera graminea (Rabenh. Ex. Schlech. Shoem.)]. Information on the combining ability for BLS resistance in Syria is not available. This study was conducted to evaluate, in 10 genetically diverse barley parents, general combining ability (GCA) and specific combining ability (SCA) effects towards the determination of the genetic basis of disease resistance and to estimate genetic variability for yield components and its modification by BLS. Ten parental genotypes varying in their reactions to BLS were crossed in a half-diallel mating design to generate 45 full-sib families. The families and the parents were inoculated with P. graminea and evaluated for resistance in replicated field tests (three inoculated and three non-inoculated plots). The parents chosen showed wide variations for resistance to BLS. Genetic component analysis showed significant effects for both GCA and SCA for resistance to BLS, suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. GCA effects were more important than SCA effects. Resistant parents exhibited high negative GCA indicating that additive gene effects were more predominant, and suggesting their prime suitability for use in barley breeding programs to improve resistance to BLS. Narrow-sense heritability was 58% and broad-sense heritability was 99% indicating that selection for BLS resistance should be effective in these crosses. A high genetic variability for the agronomic traits studied was observed. Yield components decreased significantly in inoculated plants and more pronounced in diseased plants. Significant GCA was observed for all traits. Values for GCA were, in some cases, significantly modified by BLS. This indicates that attention must be paid to the danger of drawing conclusion in quantitative genetics studies dealing with both diseased and healthy plants. Two genotypes, Banteng and Igri, had high negative GCA effects and are promising parents for enhancement of BLS resistance.     

Breeding for resistance to fusarium oxysporumF. SP. tulipae in tulip (tulipaL.).2. phenotypic and genotypic evaluation of cultivars

Summary Fusarium bulb rot is a serious tulip disease. Breeding for resistance may contribute considerably to a solution of the problem.It has been demonstrated that juvenile and adult bulbs of the same cultivars in Fusarium contaminated soil showed good agreement in degree of resistance.From an incomplete diallel cross of these cultivars second-year bulblets of 62 progenies were planted in both contaminated and non-contaminated soil. The percentages of non-diseased bulbs harvested provided a criterion for resistance. The analysis of combining ability for the degree of resistance revealed that both the mean square of GCA and that of SCA were significant. The relative magnitudes of the GCA and SCA mean squares suggest that resistance is governed primarily by additive gene action. The GCA of individual parents could be estimated and tested. In general it corresponded with their phenotypic behaviour.     

Diallel analysis of sweetpotatoes for resistance to sweetpotato virus disease

Sweetpotato virus disease (SPVD) is due to the dual infection and synergistic interaction of Sweetpotato feathery mottle potyvirus (SPFMV) and Sweetpotato chlorotic stunt crinivirus(SPCSV), and causes up to 98% yield loss in sweetpotato in East Africa. This study was conducted to determine the inheritance of resistance to SPVD in sweetpotato and to estimate the nature of genetic variance. Ten parental clones varying in reaction to SPVD were crossed in a half diallel mating design to generate 45 full-sib families. The families were graft-inoculated with SPCSV and SPFMV to induce SPVD and evaluated for resistance in a randomized complete block design at two sites in Namulonge, Uganda during 1998–2000. In serological assays for SPFMV and SPCSV,resistance to symptom development and recovery from initial systemic SPVD symptoms, characterised resistant genotypes. Genetic component analysis showed significant effects for both general combining ability (GCA) and specific combining ability (SCA) for resistance to SPVD. GCA to SCA variance component ratios were large (0.51–0.87), hence GCA effects were more important than SCA effects. Resistant parents exhibited high GCA indicating that additive gene effects were predominant in the inheritance of resistance to SPVD and recovery. Narrow-sense heritability (31–41%) and broad-sense heritability (73–98%) were moderate to high, indicating that rapid genetic gains for SPVD resistance could be accomplished by mass selection breeding techniques. Two genotypes, New Kawogo and Sowola, had high negative GCA effects and had several families in specific crosses,which exhibited rapid recovery from SPVD,and are promising parents for enhancement of SPVD resistance and recovery. This revised version was published online in August 2006 with corrections to the Cover Date.     

Combining ability analysis of resistance to head blight caused by <Emphasis Type="Italic">Fusarium graminearum</Emphasis> in spring wheat

Fusarium head blight (FHB) caused by Fusarium spp. is one of the most important fungal diseases of wheat (Triticum aestivum L.) in regions with wet climatic conditions. Improvement of the FHB resistance by developing new varieties requires sound knowledge on the inheritance of resistance. An 8 × 8 diallel analysis was performed to estimate general (GCA) and specific (SCA) combining ability of resistance to FHB. The F₁s and parental lines were evaluated under artificial inoculation at the experimental field of IFA-Tulln, Austria during 2001 and 2002. Disease severity was evaluated by repeated scoring of the percentage of infected spikelets and calculating an area under the disease progress curve (AUDPC). The analysis of combining ability across two years showed highly significant GCA and non-significant SCA effects indicating the importance of additive genetic components in controlling FHB resistance. The significant GCA-by-year interaction presented the role of environmental factors in influencing the FHB reaction of wheat lines. The comparison of the crosses with low FHB infection and GCA effects of their parents showed that such crosses involved at least one parent with high or average negative GCA effect. The results revealed that it is feasible to use highly or moderately resistant genotypes and conventional breeding methods to achieve genetic improvement of FHB resistance in spring wheat.     

Genetics of resistance to 3 isolates of Ascochyta fabae on Faba bean (Vicia faba L.) in controlled conditions

Summary Genetics of resistance to Ascochyta fabae Speg. in Vicia faba L. was studied with a final objective to develop resistant faba bean varieties to Ascochyta blight. The study was conducted separately on 3 single spore isolates (AF10-2 and AF13-2 from Tunisia and AF4-3 from France) and belonging to different groups of virulence (GV1 and GV2). Important general combining ability (GCA) effects were found especially with isolates AF10-2 and AF4-3. Specific combining ability (SCA), although significant for the 3 isolates, was important only with AF13 -2, but less important than GCA. Additive gene effects were predominant to non-additive effects. Lines 29H and A8817 transmitted to their progenies resistance to the 3 isolates, whereas 14–12 and 19TB conferred resistance to their progenies only with isolates AF13-2 and AF4-3, respectively. In the material studied, resistance was generally controlled by dominant genes but also could be attributed to recessive genes although less frequent. Analysis of segregation in the F2 of 2 crosses between the resistant lines (A8817 and 29H) and the susceptible line (14–12) with isolate AF4-3 revealed dominant monogenic control at the level of leaves in the 2 resistant lines and, in addition, a recessive gene controlling resistance of stems. Non-allelic interactions were occasionally manifested and their origin appeared to be due to line 19TB. A recurrent selection scheme was proposed with the objective to develop improved open-pollination populations and synthetic varieties responding to the objective of the national Tunisian research programme on faba bean.     

Estimates of combining ability for resistance of winter rye to Fusarium culmorum head blight

Summary Head blight caused by Fusarium culmorum and F. graminearum is damaging in all winter rye (Secale cereale L.) growing areas. For hybrid breeding, the relative magnitude of general (GCA) and specific combining ability (SCA) is a crucial parameter for developing appropriate selection procedures. Forty single-cross hybrids were produced by crossing six and seven inbred lines of the Petkus and Carsten gene pool, respectively, in a factorial design. Hybrids were evaluated in two years with artificial F. culmorum inoculation. Resistance traits were head blight rating and grain weight relative to the non-inoculated control. Both resistance traits were closely correlated across both years (r-0.8, P=0.01). Significant genotypic variation was found for both traits with medium to high estimates of heritability (h²=0.6-0.8). Components of variance for GCA were, across years, 10 and 6 times larger than those for SCA for head blight rating and relative grain weight, respectively. Significant SCA effects were found for 15 to 20% of all cross combinations across both traits in each year. SCA effects were, however, inconsistent over years leading to a high SCA-year interaction. In conclusion, resistance to Fusarium head blight among the interpool hybrids tested was conditioned mainly by additive gene action that could be utilized by recurrent selection in multi-environment trials.Abbreviations GCA general combining ability - SCA specific combining ability     

Combining ability analysis for harvest index in winter wheat

Summary Cereal breeders have used harvest index (HI) as a selection criterion in segregating generations to identify physiologically superior lines with improved partitioning of total assimilate into grain. Information on combining ability for HI of the hard red winter wheat (Triticum aestivum L.) cultivars of the Southern Great Plains is not available. A study was undertaken to examine HI of seven genetically diverse winter wheat parents, evaluate their general combining ability (GCA) and specific combining ability (SCA) effects, and study correlations of HI with several agronomic traits. The seven parents were crossed in a half-diallel mating design to produce 21 crosses. The F₁'s, their F₂ progenies, and the parents were evaluated in replicated field tests at Stillwater and at Lahoma, OK. The combining ability analysis was performed using Griffing's Method 4, Model 1. The results showed significant variation among parents for HI. The GCA and the SCA effects were mostly inconsistent between generations and between environments. However, parents with consistently high HI and positive GCA estimates were identified. The progeny with high HI mostly resulted from parents with high GCA estimates. The correlations between HI and agronomic traits indicated that improvement in HI should also result in high grain yield, early maturity, and short plant height.     

Diallel analysis of resistance to Septoria tritici isolates in durum wheat

Summary All crosses, except for reciprocals, were made among ten cultivars originating from crop improvement programs in North Africa and the Middle East. The entries varied widely in reaction to Septoria tritici. F₁ and F₂ progenies of the crosses were evaluated using eight S. tritici isolates from seven countries in the Mediterranean area. Thus, sixteen separate combining ability analyses were excecuted. General combining ability (GCA) was the major component of variation, although specific combining ability (SCA) was present in most cases. Additive variance thus appears to be of predominant importance. Nevertheless, non-additive variance may interfere when line selection in a breeding program is practiced. While differing greatly among cultivars, specific GCA effects for each cultivar separately were of similar magnitude for all isolates. Ranking statistics determined that cultivars were ranked in similar order for both means and specific GCA effects independent of the isolate used. Different isolates may therefore interact with similar or identical genetically controlled mechanisms in a particular cultivar. This could indicate the absence of differential gene-for-gene relationships and suggests that isolates vary in aggressiveness rather than in virulence.     

Resistance to Coffee Berry Disease in Arabica Coffee Variety 'Ruiru 11'

General and specific combining ability effects for resistance to coffee berry disease (Colletotrichum coffeanum Noack sensu Hindorf) in Coffea arabica variety ‘Ruiru 11’ and its parents were estimated according to the North Carolina Design II. A set of nine male parents and eleven females were crossed in all possible combinations to generate ninety-nine single cross hybrids constituting the ‘Ruiru 11’ variety. The parents and their hybrid progenies were screened and the mean score data analyzed for possible genetic variation. The contributions of the parents to the total genetic variance were low hence the low narrow sense heritability (h²= 0.04). The male and female parents accounted for 9.36 % and 0.96 %, respectively, while their interaction effect accounted for 89.68 %. In the analysis of combining ability for resistance, ExB3.96 and Cat.90 exhibited significant negative general combining ability (GCA). Specific combining abilities (SCA) were negative and significant for Cat.86 × ExB3.99, Cat.127 × ExB3.116 and Cat.119 × ExB3.879 crosses. The implications of these results in selection for highly resistant ‘Ruiru 11’ progenies are discussed.     

Top‐cross vs. poly‐cross as alternative to test‐cross for estimating the general combining ability in potato

Test‐crosses for estimating combining ability in potato although useful are difficult to make being labour intensive and time consuming, and due to the problem of sterility. The objective of the present study was to test methods for estimating general combining ability (GCA) of parents based on fewer matings. The GCA of 12 females estimated based on matings with specific single testers (top‐cross) and three types of bulk pollen (poly‐cross) were compared with those estimated by 72 (12 × 6) test‐crosses, using common testers in three types of matings. The study was conducted for 2 years at each of the two locations representing short‐days subtropical conditions in plains and long‐days temperate conditions in hills of India. Analysis of variance of test‐crosses showed that both general and specific combining ability effects were important for tuber yield, late blight resistance as well as general impression, the characters studied. Correlation coefficients among three types of matings showed that bulk pollen matings had poor effectiveness in predicting the GCA of the females, whereas matings with selected individual testers resulted in GCA estimates similar to those based on six testers. Thus use of top‐cross involving selected tester could substantially reduce the number of matings required for estimating the GCA. Among the six testers used, the tester CFK69.1 could estimate the GCA of the females with reasonably high accuracy both for tuber yield and general impression, and ‘Tobique’ for reaction to late blight.     

Diallel analysis of partial resistance to powdery mildew caused by Erysiphe graminis f. sp. hordei in spring barley (Hordeum vulgare L.)

Summary Six spring barley genotypes with growth stage dependent expression of partial resistance to powdery mildew were crossed following a half diallel scheme. Data for percentage infected leaf area at different growth stages from the F1 and F2 generation and the F2 offspring were analyzed. The absence of a contrast between parents and offspring indicated that dominance effects were not important at the young growth stages. Diallel analysis showed that general combining ability effects (GCA) were important at all growth stages. Although significant, specific combining ability effects were of minor importance. The cross between the most resistant parents with the largest, negative GCA resulted in the most resistant progeny.     

Correlation between late blight resistance and foliage maturity type in potato

The genetics of race-non-specific foliage resistance against Phytophthora infestans, of foliage maturity type, and of their association in potato (Solanum tuberosum) were studied. Six progenies were derived from a half-diallel set of crosses between diploid potato clones that represented a broad pool within the genus Solanum and were free of any of the 11 known R genes for late blight resistance. The progenies were evaluated for resistance to late blight and for foliage maturity type, and five of them showed a significant correlation between the two traits. The correlation did not account for all variation that was present for both traits, as reflected in the analysis in which the relative AUDPC values were adjusted for foliage maturity type. The present study adds to previous results: resistance against P. infestans always coincides with late foliage maturity. However, the results also indicate that some selection for late blight resistance without affecting the foliage maturity type should be possible.     

Inheritance of early blight resistance in diploid potatoes

Early blight (Alternatia solani) is a fungal disease in hot and humid environments, which causes leaf, stem and tuber lesions. Early blight resistance should be incorporated into potato cultivars because the fungicide spraying is an expensive solution for developing countries. The diploid cultivated species Solanum tuberosum group Phureja and group Stenotomum are sources of resistance alleles. The elucidation of the inheritance for early blight resistance must help to decide what could be the best breeding procedure to improve this diploid germplasm and transfer the resistance to the tetraploid level. Three experiments were carried out under controlled and field conditions to determine the heritability of this trait using nested and diallel mating designs with haploid, species and haploid-species hybrids. The narrow-sense heritability estimates were relatively high (0.64–0.78). This means that additivity was the most important type of gene action for determining resistance to early blight at the diploid level. The results suggested that diploid parents showing highest levels of resistance, throughout the cycle of disease development, can be used in 4x×2x crosses to obtain resistant tetraploid progenies to this fungal disease.     

Half-sib progeny evaluation and selection of potatoes resistant to the US8 genotype of Phytophthora infestans from crosses between resistant and susceptible parents

The objectives of this study were to evaluate the use of potato (Solanum tuberosum L.) late blight (Phytophthora infestans (Mont.) de Bary) resistant parents in cultivar development and identify superior clones possessing moderate to high late blight resistance combined with acceptable maturity and tuber quality. Ninety-five crosses were made between eight unadapted parents with reported late blight resistance (B0718-3, Bertita, Bzura, Greta, Libertas, Stobrawa, Tollocan and Zarevo) and susceptible parents (cultivars or advanced breeding clones) adapted to North American growing conditions. A total of 408 field selected clones were assessed for late blight resistance in the greenhouse and in the field using a mixture of US8 P. infestans isolates (A2 mating type, metalaxyl resistant) that overcame all known R-genes except R8 and R9. Clones with ≤ 10% infected foliar area in the greenhouse test or ≤ 0.30 RAUDPC (relative area under the disease progress curve) value in the field in 1998 were re-tested in 1999. A total of 118 (29% of 408) putative late blight resistant clones were selected. The eight late blight resistant parents differed in both the ability to transmit late blight resistance and in the level of resistance transmitted to the progeny. The Tollocan and B0718-3 families (half-sib progeny) had the greatest degree of resistance and frequency of resistant clones. Scott-Knott cluster analysis ranked 79 clones (67% of 118) in the high and moderate late blight resistant groups. Among these 79 clones, 19 clones had vine maturity equal to or earlier than mid-season combined with acceptable tuber quality. Further selection in 2000 resulted in eight advanced selected clones (six from Tollocan and two from B0718-3 families) with the same level of resistance as the parent combined with vine maturity and tuber quality equivalent to Atlantic, a standard cultivar for chip processing in North America. The results indicate that this breeding approach can be used to select parents for late blight resistance breeding and to identify superior clones with high levels of late blight resistance and marketable vine maturity and tuber quality. This revised version was published online in July 2006 with corrections to the Cover Date.     

Analysis of diallel crosses between wheat genotypes with genetically different resistance to Pseudocercosporella herpotrichoides (Fron) Deighton

Two diallels were analysed for general combining ability (GCA) and specific combining ability (SCA) to study the resistance of crosses‐between wheat genotypes, advanced to the F₅ generation, to Pseudocer‐cosporella herpotrichoides. The parents either carried the resistance‐gene Pch‐1 or had different levels of quantitative resistance, one genotype was susceptible. At medium milk‐ripening, significant effects were‐found for GCA and SCA. GCA effects were the more important. Diallel crosses between genotypes, all carrying Pch‐1, revealed interactions‐of the gene with the genotypic background. Some combinations had a‐higher level of resistance than the best parent. In these populations'CH‐75417’ was involved as a parent. Both ‘CH‐75417’ and ‘F–210.13.4.42’ had significant GCA effects. Crosses between quantitatively resistant parents yielded populations that transgressed both parents. The increased resistance level was associated with ‘Cappelle‐Desprez’, distinguished by its high GCA. In some crosses SCA contributed significantly to an increase in resistance level. Selection for resistance within the best advanced populations is recommended since it‐takes advantage of additive gene action and the high heritability estimates based on ELISA values in plant progenies.     

Analysis of diallel cross for some physical-quality traits in peanut (Arachis hypogaea L.)

The combining abilities for physical-quality traits in peanut were examined to understand the type of gene action governing these traits, and to identify peanut genotypes suitable for use as parents in breeding for quality improvement. The F₁ hybrids including reciprocals from a six-parent diallel cross along with the parents were evaluated in a randomised complete block design. Data were recorded on five quality traits in peanut viz., shelling outturn, 100-pod weight, 100-seed weight, Count and proportion of sound mature seeds. Substantial genetic variability was observed among the hybrids for the traits studied. Diallel analysis indicated that expression of majority of the quality traits is regulated predominantly by additive gene action suggesting possibility of early-generation selection, while non-additive gene action also plays an equally important role in the control of seed size. Significant reciprocal effect for all the traits denoted role of maternal parent in the expression of quality traits and importance of parental selection in quality breeding. Genotypes ICGV 86564 and TPG 41 were good combiners for seed size, while J 11 was a good combiner for improvement of shelling outturn and proportion of mature seeds. Association between general combining ability (GCA) effects and mean performance suggested that the performance per se of the genotype should be a good indicator of its ability to transmit the desirable quality attributes to its progenies. Though performance of crosses was found to be independent of parental GCA status, it is evident that at least one of the parents used in hybridisation should have large pods and seeds for obtaining better segregants.     

Combining ability analysis of Fusarium head blight resistance in western European wheat lines

The inheritance of Fusarium head blight (FHB) resistance was investigated in eight western European wheat lines using a half-diallel of F₁ crosses. The parents and F₁ crosses were point-inoculated, with a highly aggressive isolate of Fusarium graminearum, in replicated field and glasshouse trials. Type II resistance was assessed by measuring the % FHB spread and % wilted tips. There was a good correlation between the two disease parameters, % FHB spread area under the disease progress curve (AUDPC) and % wilted tips AUDPC (r = 0.86, P < 0.01). Correlation coefficients between the field and glasshouse environments were r = 0.46 (P < 0.01) for % FHB spread AUDPC and r = 0.40 (P < 0.05) for % wilted tips AUDPC. Both general combining ability (GCA) and specific combining ability (SCA) effects influenced the inheritance of FHB resistance, suggesting that in this set of parents both additive and non-additive (dominance or epistatic) effects influence the inheritance of type II FHB resistance. Highly significant GCA-by-environment (P < 0.0001) and SCA-by-environment (P < 0.005) interactions were also observed. Specific combinations of western European wheat varieties were identified with type II FHB resistance at a level equal to or more resistant than the winter wheat variety ‘Arina’.