Inheritance of resistance to two Septoria tritici isolates in spring and winter bread wheat cultivars

Summary All possible crosses (including reciprocals) were made among four winter bread (Aurora, Bezostaya 1, Kavkaz, and Trakia) and two Israeli spring wheat cultivars (spring x winter diallel), and among two South American spring wheats (Colotana and Klein Titan) with the same Israeli cultivars (spring x spring diallel) to study the inheritance of resistance to septoria tritici blotch. Parents, F₁, F₂ and backcrosses were grown in two separated blocks in the field over two years. One block was inoculated with isolate ISR398A1 and another with ISR8036. Each plant was assessed for plant height (cm), days to heading (from emergence or transplanting), and percent pycnidia coverage on the four uppermost leaves. Plant height and maturity had insignificant effects on pycnidia coverage. No cytoplasmic effects could be detected. In the spring x winter diallel general combining ability (GCA) was the major component of variation. Significant specific combining ability (SCA) was present in all cases. Partial dominance was operative in populations inoculated with ISR398A1. Resistance in the winter wheats was controlled by a small number of genes (usually two). The four winter wheats derive their resistance to ISR398A1 from their common parent Bezostaya 1 which lacks the 1B/1R wheat-rye translocation. Their resistance is readily overcome by ISR8036. Inheritance of the South American wheats can be explained by additive effects, with a small number of genes of recessive mode affecting resistance to both isolates. Breeding strategies that favor additive, and additive x dominance gene action should be pursued.     

Genetic variation for resistance to Fusarium head blight in bread wheat

Summary During a four year period, a total of 258 winter and spring wheat genotypes were evaluated for resistance to head blight after inoculation with Fusarium culmorum strain IPO 39-01. It was concluded that genetic variation for resistance is very large. Spring wheat genotypes which had been reported to be resistant to head blight caused by Fusarium graminearum were also resistant to F. culmorum. The resistant germplasm was divided into three gene pools: winter wheats from Eastern Europe, spring wheats from China/Japan and spring wheats from Brazil. In 32 winter wheat genotypes in 1987, and 54 winter wheat genotypes in 1989, the percentage yield reduction depended on the square root of percentage head blight with an average regression coefficient of 6.6. Heritability estimates indicated that for selection for Fusarium head blight resistance, visually assessed head blight was a better selection criterion than yield reduction.     

Inheritance of resistance to the chlorosis component of tan spot of wheat caused by Pyrenophora tritici-repentis, races 1 and 3

The fungus Pyrenophora tritici-repentis, causal agent of tan spot of wheat, produces two phenotypically distinct symptoms, tan necrosis and extensive chlorosis. The inheritance of resistance to chlorosis induced by P. tritici-repentis races 1 and 3 was studied in crosses between common wheat resistant genotypes Erik, Hadden, Red Chief, Glenlea, and 86ISMN 2137 and susceptible genotype 6B-365. Plants were inoculated under controlled environmental conditions at the two-leaf stage and disease rating was based on presence or absence of chlorosis. In all the resistant × susceptible crosses, F₁ plants were resistant and the segregation of the F₂ generation and F₃ families indicated that a single dominant gene controlled resistance. Lack of segregation in a partial diallel series of crosses among the resistant genotypes tested with race 3␣indicated that the resistant genotypes possessed␣the same resistance gene. This resistance gene was effective against chlorosis induced by P.␣tritici-repentis races 1 and 3.     

Inheritance and allelism of Russian wheat aphid resistance in several wheat lines

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko) has caused serious reduction in wheat production in 17 Western states of the United States since 1986. Inheritance of resistance to RWA in seven wheat lines and the allelism of the resistance genes in these lines with three known resistance genes Dn4, Dn5, and Dn6 were studied. The seven resistant lines were crossed to a susceptible wheat cultivar ‘Carson’ and three resistant wheats: CORWA1 (Dn4), PI 294994 (Dn5), and PI 243781 (Dn6). Seedlings of the parents, F₁, and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21–28 days after the infestation using a 1–9 scale. The resistance level of all the F₁ hybrids was similar to that of the resistant parent, indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ population, suggesting qualitative, nonadditive gene action, in which the presence of any one of the dominant alleles confers complete resistance to RWA. Resistance in CI 2401 is controlled by two dominant genes. Resistance in CI 6501 and PI 94365 is governed by one dominant gene. Resistance in PI 94355 and PI 151918 may be conditioned by either one dominant gene or one dominant and one recessive gene. No conclusion can be made on how many resistance genes are in AUSVA1-F₃, since the parent population was not a pure line. Allelic analyses showed that one of resistance genes in CI 2401 and PI 151918 was the same allele as Dn4, the resistance gene in CI 6501 was the same allele as Dn6, and AUS-VA1-F₃ had one resistance gene which was the same allele as one of the resistance genes in PI 294994. One non-allelic resistance gene different from the Dn4, Dn5, and Dn6 genes in CI 2401, PI 94355, PI 94365, and PI 222668 was identified and should be very useful in diversifying gene sources in wheat breeding.     

Inheritance and allelism of resistances to the Russian wheat aphid in seven wheat lines

Summary Studies were conducted to determine the inheritance and allelic relationships of genes controlling resistance to the Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), in seven wheat germplasm lines previously identified as resistant to RWA. The seven resistant lines were crossed to a susceptible wheat cultivar Carson, and three resistant wheats, CORWA1, PI294994 and PI243781, lines carrying the resistance genes Dn4, Dn5 and Dn6, respectively. Seedlings of the parents, F₁ and F₂ were screened for RWA resistance in the greenhouse by artificial infestation. Seedling reactions were evaluated 21 to 28 days after the infestation using a 1 to 9 scale. All the F₁ hybrids had equal or near equal levels of resistance to the resistant parent indicating dominant gene control. Only two distinctive classes were present and no intermediate types were observed in the F₂ segregation suggesting major gene actions. The resistance in PI225262 was controlled by two dominant genes. Resistance in all other lines was controlled by a single dominant gene. KS92WGRC24 appeared to have the same resistance gene as PI243781 and STARS-9302W-sib had a common allele with PI294994. The other lines had genes different from the three known genes.     

Influence of the D genome in conferring resistance to fusarium head blight in spring wheat

Reciprocal crosses were made between resistant hexaploid spring wheat cultivars/lines Sumai 3, Ning8331, and 93FHB21, and susceptible tetraploids Stewart 63 and DT486 to generate 35 chromosome pentaploids. Four heads from each of five F₁ pentaploid plants from each cross were screened with Fusarium graminearum for fusarium head blight (FHB) reaction. No pentaploid was as resistant to FHB as the resistant parents. Pentaploids derived from several crosses were more resistant than the susceptible parents, a few were more susceptible, and all plants from crosses with 93FHB21 failed to survive. Most viable seeds were obtained from the cross Sumai 3 × DT486. From this cross four of the five F₁ pentaploid parents were fertile and 354F₂ seeds derived from these four pentaploids were sown and evaluated for their FHB reaction. The majority of F₂ plants from pentaploids 1 and 3had the visual appearance and level of resistance of Sumai 3, whereas progeny from pentaploids 4 and 5 were more varied morphologically and generally more susceptible. Forty-three of the screened F₂ plants were tested for the presence of specific D chromosomes by wheat microsatellite analysis. There was no relationship between presence/absence of D chromosomes and FHB reaction. Twenty-four lines had all D chromosomes present of which 10 were intermediate-susceptible and 14 were resistant to FHB. Three lines, one resistant and two intermediate, had no D chromosomes. The remainder had between 1 and 6 of the D chromosomes present and ranged from resistant to susceptible in FHB reaction. It appears that FHB resistance is not conferred by the D genome of Sumai 3. This revised version was published online in July 2006 with corrections to the Cover Date.     

Monosomic analysis of Helminthosporium leaf blight resistance genes in wheat

A set of 21 monosomic (2n ‐ 1) and the disomic (2n) lines of the ‘Chinese Spring’ cultivar were crossed with ‘Chirya‐3′, the CIMMYT synthetic wheat line which has been identified as highly resistant for Helminthosporium leaf blight disease (HLB), in order to locate the genes governing disease resistance. The F₁ and segregating populations were challenged and screened against the most virulent pure mono‐conidial HLB isolate KL‐8 (Karnal, India). The F₁ progenies of the crosses were found to be susceptible because of the recessive nature of resistance. The F₂ progeny of the control cross (‘Chinese Spring’בChirya‐3’), segregated in the ratio of 1: 15 (resistant: susceptible), indicating that resistance to HLB was controlled by a pair of recessive genes. While the F₂ progeny of 19 monosomic crosses segregated in the ratio of 1: 15 (resistant: susceptible), the progeny of the remaining two crosses, 7B and 7D, deviated significantly from the ratio, revealing that 7B and 7D were the critical chromosomes for resistance genes that were located one on each chromosome. Moreover, the critical lines, 7B and 7D, confirmed the digenic complementary recessive nature of gene action by fitting well with the overall pooled F₂ segregation ratio of 13: 51 (resistant: susceptible) as expected for digenic complementary recessive resistance. The F₃ segregation ratios of the critical crosses, based on their pooled F₂ analysis, was estimated as 19: 32: 13 (non‐segregating susceptible: segregating as susceptible and resistant: non‐segregating resistant). F₃ progenies when tested with these ratios showed goodness‐of‐fit, confirming that the two pairs of recessive resistance genes were located on chromosomes 7B and 7D.     

Recessive genes controlling resistance to Helminthosporium leaf blight in synthetic hexaploid wheat

A study was conducted under controlled environment conditions in a phytotron to determine the nature of the inheritance of resistance Helminthosporium leaf blight (HLB) in a synthetic hexaploid wheat line, ‘Chirya‐3’, against the isolate KL‐8 of Bipolaris sorokiniana from the major wheat growing region of India. Crosses were made between two susceptible lines ‘WH 147’ and ‘Chinese Spring’. Analyses of F₁ and F₂ populations of these two crosses (‘WH 147’בChirya‐3’ and ‘Chinese Spring’בChirya‐3’) showed that resistance against the isolate in ‘Chirya‐3’ was governed by two recessive genes functioning in a complementary interaction giving an F₂ segregation pattern of 1 : 15 (resistant : susceptible). The segregation pattern of the resistant F₂ progenies in F₃ families from both crosses confirmed that two homozygous recessive genes were responsible for resistance to the isolate of Bipolaris sorokiniana in the synthetic line ‘Chirya‐3’. It is proposed that the genes be designated as hlbr1 and hlbr2.     

Influence of winter and spring wheat genetic backgrounds on haploid induction parameters and trait correlations in the wheat × maize system

The interactive influence of winter and/or spring wheat genetic background on haploid induction parameters and trait correlation was studied by hybridizing five elite and diverse genotypes each of winter and spring wheat and their F₁s (winter × winter, spring × spring, and winter × spring, generated in a diallel design excluding reciprocals) with a single genotype of maize. Data were recorded with respect to per cent seed formation, embryo formation, and regeneration. High genetic variability was present among the wheat genotypes (parents + F₁s) for the three haploid induction parameters. Significant differences were obtained within and between different groups viz., spring wheats, winter wheats, spring × spring wheats, winter × winter wheats, and winter × spring wheats with respect to the three haploid induction parameters based on ANOVA. The winter genotypes (winter parents and winter × winter wheat hybrids) responded better than the spring groups (spring wheat parents, spring × spring and winter × spring wheat hybrids) with respect to embryo formation and winter × spring wheat hybrids yielded significantly the highest numbers of regenerants. Correlation studies amongst the haploid induction parameters indicated that the genes controlling seed formation and haploid plantlet regeneration are negatively correlated when the genetic backgrounds of both ecotypes are combined in winter × spring hybrids. Haploid embryo formation had no association with seed formation and regeneration in all genetic backgrounds, suggesting independent inheritance.     

Distribution and allelic expressivity of genes for hybrid necrosis in some elite winter and spring wheat ecotypes

The distribution and allelic expressivity of hybrid necrosis genes (Ne ₁ and Ne ₂) were studied in 21 winter (mostly exotic) and 43 spring type elite wheat genotypes, by crossing them with two known testers, C 306 (Ne ₁-carrier) and HD 2380 (Ne ₂-carrier).Ne ₁ gene was present in one north-west Himalayan winter wheat landrace, Shoure Local, but absent in the other winter as well as spring wheats. Ne ₂ gene was prevalent to a much lower extent in the exotic winter wheat germplasm (31.57%) as compared to the recently developed Indian and Mexican spring wheat semidwarfs (69.80%). This may suggest that breeders have tried to preclude hybrid necrosis by selecting for non-carrier genotypes in the development of exotic winter wheats in contrast to the situation in spring wheats. Based on the degree of expression of hybrid necrosis genes in the F₁ hybrids, the carrier genotypes were characterized with respect to the allelic strength of the hybrid necrosis genes. The 27 non-carrier genotypes of the two ecotypes identified in the present study have a greater potential use in future hybridization programmes so as to overcome the problem of hybrid necrosis. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance and allelism of wheat streak mosaic virus resistance in two Iranian wheat lines

Wheat streak mosaic (WSM) caused by wheat streak mosaic virus (WSMV) is a serious disease of wheat and many plants in the grass family. In previous studies genotypes collected from different parts of Iran, were screened for WSM resistance. Two resistant genotypes, “Adl Cross” and “4004” were crossed to one susceptible genotype “Marvdasht.” Reciprocal crosses were also made. Seedlings of parents, F₁, F₂, backcrosses to susceptible (BCS) and resistant (BCR) were evaluated for WSMV reaction based on scales 0–7, by artificial infection under greenhouse conditions. Allelism was studied by evaluating the F₁ and F₂ seedlings of the resistant × resistant cross. Inheritance of resistance to WSMV in Adl Cross and 4004 was controlled by one dominant gene. No allelism was observed between resistance genes. Among the F₂ seedlings of the resistant × resistant cross relatively more resistant genotypes with a zero score were observed. These two genes, therefore, can be incorporated into an adapted wheat cultivar to produce a more durable resistance.     

Winter × spring wheat hybridization - a promising avenue for yield enhancement

Introgression of the winter gene pool into spring wheat is being considered as one of the strategies to break through the yield plateau. However, little information is available on the combining ability of these two important but distinct groups of wheats in Indian conditions. Therefore, the present study was undertaken to determine the combining ability and gene action of yield and yield attributes in winter × spring wheat crosses. Seventy F₁ progenies developed by 14 winter and five spring wheat lines using a line × tester design were evaluated, along with their parents, for yield and yield attributes in a randomized complete block design under field conditions. The mean squares for all the characters studied showed highly significant differences. The mean squares due to female × male interactions were significant for all the characters studied except for grains per ear and grain weight per ear. Additive genetic effects were found to play a key role in controlling the expression of days to heading, plant height and spikelets per ear.‘MV 19’ and ‘Stepniak’/‘Karvuna’ among winter and ‘PBW 65’ among spring wheats were good general combiners for most of the yield attributes studied. The estimates for specific combining ability effects suggested that, although general combining ability (GCA) effects of most winter wheats are either average or poor, their combination can give desirable genotypes with spring wheat parents possessing a high GCA.     

The inheritance of resistance to beet necrotic yellow vein virus (BNYVV) in B. vulgaris subsp. maritima, accession WB42: Statistical comparisons with Holly-1-4

In this study, the inheritance of resistance to Beet necrotic yellow vein virus (BNYVV) in accessions Holly-1-4and WB42 was investigated. Crosses between both resistant sources and susceptible parents were carried out and F₁F₂ and BC₁ populations were obtained. Virus concentrations in WB42and its F₁ populations were lower than in Holly-1-4. Observed ratios of susceptible and resistant plants in segregating populations of Holly-1-4 as well as WB42 were in agreement with hypothesis of one dominant major gene. Segregation of plants in F₂ populations obtained from crosses betweenHolly-1-4 and WB42 revealed that the resistance genes in Holly-1-4 and WB42 were nonallelic and linked loci. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of Russian wheat aphid resistance in PI 140207 spring wheat

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), has become a serious, perennial pest of wheat (Triticum aestivum L.) in many areas of the world. This study was initiated to determine the inheritance of RWA resistance in PI 140207 (a RWA-resistant spring wheat) and to determine its allelic relationship with a previously reported RWA resistance gene. Crosses were made between PI 140207 and ‘Pavon’ (a RWA-susceptible spring wheat). Genetic analysis was performed on the parents, F₁, F₂, backcross (BC) population and F₂-derived F₃ families. Analyses of segregation patterns of plants in the F₁, F₂, and BC populations, and F₂-derived F₃ families indicated single dominant gene control of RWA resistance in PI 140207. Results of the allelism test indicated that the resistance gene in PI 140207, while conferring distinctly different seedling reactions to RWA feeding, is the same as Dn 1, the resistance gene in PI 137739.     

Inheritance of insensitivity to culture filtrate of Pyrenophora tritici-repentis, race 2, in wheat (Triticum aestivum L.)

Tan spot of wheat is caused by the fungus Pyrenophora tritici‐repentis. On susceptible hosts, P. tritici‐repentis induces two phenotypically distinct symptoms, tan necrosis and chlorosis. This fungus produces several toxins that induce tan necrosis and chlorosis symptoms in susceptible cultivars. The objectives of this study were to determine the inheritance of insensitivity to necrosis‐inducing culture filtrate of P. tritici‐repentis, race 2, and to establish the relationship between the host reaction to culture filtrate and spore inoculation with respect to the necrosis component. The F₁, F₂, and BC₁F₁ plants and F_2:8 lines of five crosses involving resistant wheat genotypes ‘Erik’, ‘Red Chief’, and line 86ISMN 2137 with susceptible cultivars ‘Glenlea’ and ‘Kenyon’ were studied. Plants were spore‐inoculated at the two‐leaf stage. Four days later, the newly emerged uninoculated third leaf was infiltrated with a culture filtrate of isolate Ptr 92–164 (race 2). Reactions to the spore inoculation and the culture filtrate were recorded 8 days after spore inoculation. The segregation observed in the F₂ and BC₁F₁ generations and the F_2:8 lines of all crosses indicated that a single recessive gene controlled insensitivity to necrosis caused by culture filtrate. This gene also controlled resistance to necrosis induced by spore inoculation.     

Genetic analysis of resistance to root-lesion nematode (Pratylenchus thornei) in wheat

The inheritance of resistance to root‐lesion nematode was investigated in five synthetic hexaploid wheat lines and two bread wheat lines using a half‐diallel design of F₁ and F₂ crosses. The combining ability of resistance genes in the synthetic hexaploid wheat lines was compared with the performance of the bread wheat line ‘GS50a’, the source of resistance to Pratylenchus thornei used in Australian wheat breeding programmes. Replicated glasshouse trials identified P. thornei resistance as polygenic and additive in gene action. General combining ability (GCA) of the parents was more important than specific combining ability (SCA) effects in the inheritance of P. thornei resistance in both F₁ and F₂ populations. The synthetic hexaploid wheat line ‘CPI133872’ was identified as the best general combiner, however, all five synthetic hexaploid wheat lines possessed better GCA than ‘GS50a’ The synthetic hexaploid wheat lines contain novel sources of P. thornei resistance that will provide alternative and more effective sources of resistance to be utilized in wheat breeding programmes.     

Genetic analysis of flowering and maturity time in high latitude spring wheat

Due to the short growing season in the high northern latitudes, the development of early maturing spring wheat (Triticum aestivum L.) cultivars is important to avoid frost damage which can lower production and quality. We investigated earliness of flowering and maturity, and some associated agronomic traits, using a set of randomly selected high northern latitude adapted spring wheat cultivars (differing in maturity) and their F₁ and F₂ crosses made in a one-way diallel mating design. The parents, and their F₁ and F₂ crosses were evaluated under field conditions over 2 years. Anthesis and maturity times were controlled by both vernalization response and earliness per se genes, mainly acting additively. Non-additive genetic effects were more important in controlling grain fill duration, grain yield and plant height. Additive × additive epistatic effects were detected for all traits studied except time to anthesis. Segregation analyses of the F₂ populations for time to anthesis indicated the presence of different vernalization response genes. Molecular genetic analyses revealed the presence of Vrn-A1 and Vrn-B1 genes in the parental cultivars. Narrow-sense heritability was medium to high (60–86%) for anthesis and maturity times but low to medium (13–55%) for grain fill duration, plant height and grain yield. Selection for early flowering/maturity in early segregating generations would be expected to result in genetic improvement towards earliness in high latitude spring wheats. Incorporation of the vernalization responsive gene Vrn-B1 in combination with vernalization non-responsive gene Vrn-A1 into spring wheats would aid in the development of early maturing cultivars with high grain yield potential for the high latitude wheat growing regions of the northern hemisphere.     

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.     

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.     

Inheritance of Dry Root Rot (Rhizoctonia bataticola) Resistance in Chickpea (Cicer arietinum)

The inheritance of resistance to dry root rot of chickpea caused by Rhizoctonia bataticola was studied. Parental F₁ and F₂ populations of two resistant and two susceptible parents, along with 49 F₁ progenies of one of the resistant × susceptible crosses were rested for their reaction to dry root rot using the blotting-paper technique. All F, plants of the resistant × susceptible crosses were resistant; the F₂ generation fitted a 3 resistant: 1 susceptible ratio indicating monogenic inheritance, with resistance dominant over susceptibility. F³ family segregation data confirmed the results. No segregation occurred among the progeny of resistant × resistant and susceptible × susceptible crosses.