Generation mean analysis to identify and partition the components of genetic resistance to Septoria nodorum in wheat

Summary Foliar symptom severity of seedlings artificially inoculated with S. nodorum were used to idenify the type of gne action controlling resistance to this pathogen in the early generations of two wheat crosses. In both crosses a resistant spring wheat cultivar was crossed to a susceptible cultivar. Reciprocal crosses were included in the analysis to determine if the cytoplasm contributed in any significant degree to the level of resistance present.Results indicated that resistance was polygenic and that it could be explaned prinerpally by additive gene effects. Some differences in reciprocal crosses were evident, but a significant role for the eytoplasm in resistance is not indicated.Cooperative investigations of the Agricultural Research Service of the U. S. Department of Agriculture and the Montana Agricultural Experiment Station. Journal Paper No. 1210.     

Components of partial resistance to Leptosphaeria nodorum among seven soft red winter wheats

Summary Seven soft red winter wheat cultivars were evaluated for partial resistance to Leptosphaeria nodorum under field conditions. The results demonstrate that resistance is available among cultivars that are adapted to the southeastern U.S. and that resistance is long lasting. Resistant cultivars had longer incubation and latent periods, slower rates of lesion development, and reduced the level of sporulation of L. nodorum. Seedlings of susceptible cultivars sustained severe disease with dew periods as short as 48 hr in greenhouse tests. One resistant cultivar sustained little damage with dew period as long as 144 hr.Latent period was shortest on the second leaf below the flag leaf (F-2 leaf) of all cultivars and longest on the flag leaf which reflected the effect of microclimate and leaf age. The range in length of latent period on the flag, F-1, and F-2 leaves of resistant cultivars was less than that for susceptible cultivars. Differences between cultivars were greatest (up to 6.8 days) for the F-2 leaf. A delay in production of inoculum on the F-2 and lower leaves of resistant cultivars should delay infection of the flag leaf and spike.The greatest differentiation among cultivars for sporulation of L. nodorum was on upper leaves at Feekes growth stage 11.2. Oasis consistently had less sporulation than other cultivars at all sampling dates and leaf positions. The rate of disease progress up the plant (disease severity) and area under the disease progress curve were also least on Oasis.There were significant correlations among components of resistance and associated components suggesting that a single or interrelated mechanisms control expression of resistance. Overall, Oasis was the most resistant cultivar followed by Coker 762 and Coker 747. There cultivars have remained resistant for ten or more years. Stacy was intermediate in resistance and was the most variable of the seven cultivars for the components tested. Holley, Omega 78, and Florida 301 were highly susceptible by all criteria tested. Plant height was not a factor in resistance. The resistant cultivars are mostly later in maturity than the susceptible cultivars. In this study, crop maturity was similar under existing environmental conditions during the two seasons when most data were collected.     

Inheritance,heritability and breeding behaviour of three sources of resistance to Septoria tritici in wheat

Summary We examined the inheritance, heritability, and breeding behaviour of resistance to Septoria tritict in the spring wheat cultivars Seabreeze, Veranopolis, and IAS-20 in crosses to the susceptible Australian spring cultivar Gamenya. Resistance in Seabreeze was higher than that in Veranopolis and IAS-20. Inheritance of resistance in Seabreeze seems to be determined by at least three recessive genes. Resistance in Veranopolis and IAS-20 might be due to a single gene. Similarity of inheritance patterns and breeding behaviour of the resistance from Veranopolis and IAS-20 and the common ancestry of these two cultivars suggest that they may carry the same gene. Standard unit heritabilities of Septoria scores, as measured by correlation of F₂ plant with F₃ family mean data, were high (57–68%). Parent offspring genetic correlations confirmed previously reported associations between resistance and lateness and resistance and tall plant height, but correlations were not sufficiently high to be a major obstacle to selection. Resistance in Veranopolis and IAS-20 were easily recovered in first backcrosses while that in Sea breeze was more difficult to recover.Contribution from the Department of Agriculture Western Australia.     

Genotypic variation for resistance to Septoria nodorum in triticale

Septoria nodorum leaf and glume blotch is an important disease of triticale (×_Triticosecale Wittm.) and can cause severe losses of grain yield in some regions. Quantitative genetic parameters for resistance were estimated for 2 years in two locations in triticale genotypes artificially inoculated with S. nodorum. The effect of infection was assessed by a visual symptom rating of flag leaves and spikes and by grain yield traits relative to an uninoculated control. The mean ratings of flag leaves and spikes, calculated from two to four ratings, were 2.6 and 3.9, respectively, with a range of six ratings for spikes and over five for flag leaves. Infection caused an 11.5% mean reduction in kernel weight per spike, which was the result of 13.2% lower 1000‐kernel weight. The number of kernels per spike and 50‐ml weight were little affected. For all relative grain yield traits, genotypic variation was small with high genotype‐environment interaction effects and thus moderate to low heritabilities. In contrast, for visual ratings genotypic variation was high, with low interaction effects leading to high heritabilities. Phenotypic correlation between flag leaf and spike ratings was low, indicating independent disease resistance mechanisms. The best association, although still moderate, was obtained between flag leaf rating and relative 1000‐kernel weight. Therefore, visual disease ratings do not satisfactorily assess the effect of Septoria infection on grain yield traits. The reduction in 1000‐kernel and possibly 50‐ml weight are good indicators, provided that multi‐environment tests are conducted.     

Resistance to Septoria nodorum blotch in several Triticum species

Summary Resistance to septoria nodorum blotch in Triticum monococcum, T. tauschii, T. timopheevii, T. dicoccum and T. durum was evaluated on plants at the three-leaf stage in greenhouse tests. A high frequency of resistant genotypes was found in T. monococcum, T. tauschii and T. timopheevii, but not in T. dicoccum and T. durum. The resistance of F₁ plants of crosses of resistant T. monococcum (PI 289599) and T. timopheevii (PI 290518) accessions with susceptible common wheat cv. Park and durum wheat cv. Wakooma, respectively, was evaluated on the basis of percentage leaf necrosis, lesion number, lesion size and incubation period. No dominance was found for long incubation period, but various dominance relationships occurred for low percentage leaf necrosis, low lesion number and small lesion size, depending on the cross. Multiple regression analysis showed that lesion number contributed more to percentage leaf necrosis than lesion size or incubation period. Resistance to septoria nodorum blotch was transferred successfully from T. timopheevii to cultivated durum wheat. Resistant BC₁F₇ lines, recovered from the T. timopheevii (PI 290518) × Wakooma cross, showed normal chromosome behaviour at meiosis (14 bivalents) and were self-fertile. However, an effective level of resistance was not recovered in lines derived from the other interspecific crosses.     

The influence of disease complexes involving Leptosphaeria (Septoria) nodorum on detection of resistance to three leaf spot diseases in wheat

Summary In controlled inoculation studies with Septoria nodorum and Pyrenophora tritici-repentis, estimates of the relative proportion of each pathogen demonstrated differences in the responses of cultivars to pathogen mixtures that were not apparent from measurements of diseased leaf areas. Under field conditions estimates of the relative proportion of S. nodorum, P. tritici-repentis and S. tritici varied between field screening locations in Western Australian but also between lines within locations. Lines with known resistance to P. tritici-repentis and S. tritici, but susceptible to S. nodorum, could not be distinguished from susceptible lines on the basis of leaf area diseased or grain weight depression when S. nodorum was present in the disease complex. Such conditions, while suitable for the selection of combined resistance to these pathogens, were unsuitable for identifying resistance to individual pathogens. As symptoms were similar, the proportion of diseased leaf area sporulating with each pathogen provided a means of measuring the variation in disease development induced on lines varying in resistance. Knowledge of the components of disease and their relative importance were essential in understanding varietal response information under mixed infections of these leaf spot pathogens.     

Chromosome and chromosomal arm locations of genes for resistance to Septoria glume blotch in wheat cultivar Cotipora

Summary Septoria glume blotch, caused by Stagonospora nodorum, is an important disease of wheat (Triticum aestivum). Separate genetic mechanisms were found to control flag leaf and spike resistance. Genes for resistance to S. nodorum were located on different chromosomes in the few wheat cultivars studied. These studies only partially agree on the chromosome locations of gene in wheat for resistance to S. nodorum, and chromosomal arm locations of such genes are not known. The objectives of this study were to determine the chromosome and chromosomal arm locations of genes that significantly influence resistance to S. nodorum in wheat cultivar Cotipora. Monosomic analysis showed that flag leaf resistance was controlled by genes on chromosomes 3A, 4A, and 3B whereas the spike resistance was controlled by genes on chromosomes 3A, 4A, 7A, and 3B (P=0.01). Additionally, genes on chromosomes 6B and 5A influenced the susceptibility of the flag leaf and spike reactions, respectively (P=0.01). Telocentric analysis showed that genes on both arms of chromosome 3A, and the long arms of chromosomes 4A and 3B were involved in the flag leaf resistance whereas genes on both arms of chromosome 4A, the short arm of chromosome 3A, and the long arm of chromosome 3B conferred spike resistance.     

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.     

QTL controlling partial resistance to Stagonospora nodorum leaf blotch in winter wheat cultivar Alba

Stagonospora nodorum blotch (SNB) is an important foliar and glume disease in cereals. Inheritance of SNB resistance in wheat appears quantitative. The development of partially resistant cultivars seems to be the only effective way to combat the pathogen. Partial resistance components like length of incubation period (INC), disease severity (DIS) and length of latent period (LAT) were evaluated on a population of doubled-haploids derived from a cross between the partially resistant cultivar Alba and the susceptible cultivar Begra. Experiments were conducted in controlled environments and the fifth leaf was examined. Molecular analyses were based on bulked segregant analyses (BSA) and screening with 240 microsatellites DNA markers. The QTL analysis revealed QTL on chromosome 6AL (designated as QSnl.ihar-6A) and putative QTL on chromosome 6D. The QSnl.ihar-6A accounted for 36% of the phenotypic variance for DIS and 14% for INC. The putative QTL accounted for 10% of the variability in INC and 8% of DIS components of SNB resistance.     

The response of field-inoculated wheat cultivars to mixtures of Septoria tritici isolates

Summary Wheat cultivars of diverse genetical background and response to Septoria tritici were inoculated during 2 years in the field with single or mixtures of isolates. Significant reductions in pycnidial coverage were recorded for mixtures of 2 or 5 isolates relative to the virulent isolate ISR8036 under the moderate 1989/1990 epidemic. The interactions between cultivars and all possible combinations among ISR398A1, USR8036 and the 1:1. mixture of the 2 isolates were highly significant. Cultivars exposed to mixtures of isolates expressed differential response in pycnidial coverage compared to the single isolate response. The coverage in the mixtures was significantly less than that of the arithmetic mean between the two isolates. Under the severe 1990/1991 epidemic pycnidial coverage on cultivars inoculated with the mixture of the same 2 isolates did not differ statistically from that of ISR8036, yet, ISR398A1 differed from ISR8036 and the isolate mixture. Losses in 1000-kernel weight for 12 wheat cultivars which were repeated during the 2-trial-years were significantly lower in the isolate mixture relative to that of ISR8036. The suppression of symptoms in isolate mixture relative to the expected expression of the most virulent component may be indicative of differential aggressiveness of isolates regardless of their virulence. The phenomenon may affect screening and selection procedures in breeding for resistance.     

Selection for biomass yield in wheat

Summary Biomass (above ground plant parts) yield may be a useful selection trait for yield improvement in wheat (Triticum aestivum L.). This study was conducted to estimate realized heritability of biomass yield and to determine the response to selection for high and low biomass yield in 8 genetically diverse populations of spring wheat under two production systems. Selections were made among the F₃ lines. Progenies of the selected lines were evaluated in replicated field tests in the F₄ generation under high fertility and low fertility production systems at Rampur, Nepal, in 1991. Fertility level had a significant effect on biomass yield, grain yield, effective tiller number, number of kernels per spike, thousand kernel weight, and harvest index. Selection in the F₃ for high and low biomass yield was effective in identifying F₄ lines with high and low biomass yield, respectively. Biomass yield differences between high and low selection groups in the F₄ generation, expressed as percent of the mean of the low selection group and averaged over the eight populations, were 53.9 and 36.5% higher than the mean of the low selection group under the high and the low fertility production systems, respectively. The corresponding figures for grain yield were 48.8 and 34.9% under the high and the low production systems, respectively. Also, selection for high biomass yield resulted in higher effective tiller number, and number of kernels per spike, but lower harvest index. Realized heritability estimates for biomass yield were greater at high fertility (range 0.49 to 0.85) than at low fertility (range 0.22 to 0.44). Biomass yield showed positive genotypic correlations with grain yield, effective tiller number, and number of kernels per spike but a negative correlation with harvest index. The results indicated that selection for high biomass yield should bring about positive improvements in biomass yield, grain yield, effective tiller number, and number of kernels per spike. The correlation between F₃ and F₄ generations suggested that biomass yield in the F₃ generation was a good predictor of biomass yield and grain yield in the F₄ generation. Selection for biomass yield in wheat should be made under the standard production system to obtain a realistic response.     

Comparison of resistance of triticale,wheat and spelt to septoria nodorum blotch at the seedling and adult plant stages

Summary On average, the cereal species studied were susceptible to septoria nodorum blotch (SNB), except for spring triticale on leaf and head and winter titicale on leaf, that appeared to be significantly more resistant, than the other ones.In all three species the SNB response of the adult plants was to a limited extent only predicted by the reaction on first leaf seedlings. In most cases it was impossible to predict the response to SNB of adult plants on the basis of seedling reaction. Correlations between the adult plant stage and the seedling stage, or detached seedling leaves, appears not to be sufficient for use in practical breeding work. A reversal of reaction to SNB was even found between the above growth stages in studied spring and winter wheat varieties.     

Selection for yield in small plots of spring wheat

Summary Four selection experiments were sown: single-row plots with single row-spacing (20 cm), single-row plots with double row spacing (40 cm), three-row plots and six-row plots both with single row-spacing (20 cm). Selection entries were mimiced by 16 different varieties or advanced breeding lines, which were also sown in a yield trial. Each experiment was laid out as a 4-times replicated randomized block design. Row length was 2 m. Alley borders and border-rows of multiple-row plots were harvested separately to evaluate the effects of different harvesting procedures on the selection efficiency. Removal of alley borders was found to be disadvantageous, since the gain in precision was more than offset by the loss in sampled area. Wide spacing of single-row plots improved the selection efficiency in comparison with normal spaced single-row plots. In multiple-row plots the selection efficiency was not improved by harvesting only the central rows.For gross plot yield (= yield of net plot + yield of alley borders) the differences in selection efficiency between the various selection plot-types were explained on the basis of the genetic variance, the environmental variance and the coefficient of genetic correlation with farm yield as determined in the yield trial.     

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.     

Reaction of tritordeum to Fusarium culmorum and Septoria nodorum

Summary Hordeum chilense is a wild barley extensively used in wide crosses in the Triticeae. It could be a valuable source of resistance to Fusarium culmorum and Septoria nodorum. Some H. chilense x Triticum spp. amphiploids, named tritordeums, were more resistant than the parental wheat line to these diseases, others were not. Average contents of ergosterol and deoxynivalenol (DON) suggested that resistance to colonization by Fusarium was the highest for Hordeum chilense, followed by tritordeum and wheat in decreasing order. In particular, the H. chilense genotypes H7 and H17 enhanced the wheat resistance to F. culmorum in its tritordeum offsprings. Resistance to S. nodorum in tritordeum was not associated with tall plant height. There is sufficient genetic variation for resistance to F. culmorum and S. nodorum among tritordeum to allow the breeding of lines combining short straw and resistance to both diseases.     

Selection for yield in wheat breeding

Summary If selection based on F₃ yield tests is to be effective, the yield tests must be successful in discriminating among yield genotypes. The available literature indicates that simple tests with limited or no replication are not very effective, although more extensive, replicated tests may be.Data from an experiment comparing F₃ yield tests with a single seed descent procedure showed that F₃ selection based on a two-replicate test with single seed descent procedure did not justify the extra work involved.     

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.     

An Improved Assessment Procedure for Breeding for Resistance to Septoria nodorum in Wheat

A yield-based assessment procedure for breeding for resistance to Septoria nodorum (SN) is presented. Artificially infected as well as fungicide-protected plots were analyzed for each genotype. Considerable increase in precision of disease scores was gained by using an assessment matrix for different plant organs and time intervals during disease development. This technique resulted in yield loss: disease attack correlations up to r = 0.85:** and raised coefficients of heritability (h²_b= broad sense heritability) with h²_b= 0.72 for % yield loss and h²_b= 0.86 for SN-attack, calculated for a 3-year replicated field experiment with 105 winter wheat cultivars.     

Components of resistance to Mycosphaerella graminicola and Phaeosphaeria nodorum in spring wheats

Summary Infection frequency (IF), latent period (LP) and sporulation rate (S) were assessed on 23 genotypes (cultivars, breeding lines and parents) in the field and 36 genotypes in the glasshouse (with 22 being in common) following inoculation with conidia of either Mycosphaerella graminicola or Phaeosphaeria nodorum. Significant variation was observed among genotypes for each resistance component. Genotypes resistant to Mycosphaerella graminicola had long LP and low S but variable IF. Genotypes resistant to Phaeosphaeria nodorum had low IF, long LP and low to moderate S. IF and S were independent while a negative correlation was observed between LP and S for Mycosphaerella graminicola only. LP was most consistently correlated with field response. Components measured in the field better discriminated resistant genotypes from others. Combinations of genotypes were sought for which one genotype may contribute to a reduced disease severity for S. nodorum and the other genotype may contribute to a reduced disease severity for S. tritici in mixtures. Such combinations of genotypes were rare.     

Selection for Yield and its Components in a Winter Wheat Population under Different Environmental Conditions in Albania

Wheat breeding programmes in Albania depend upon the use of diverse genetic material of local origin and on special environmental conditions; the present study being conducted under drought stress conditions. Grain yield, yield components, plant height and test weight were estimated in an F₅ generation of a winter wheat population at Kamez in 1989—90 and 1990—91 and at Lushnje in 1989—90. The highest values of heritability were observed for plant height, kernel weight and spike length. Due to drought stress the heritability of kernels per spike was low. None of the correlated responses was higher than the direct response for yield. Maximum genetic gain was expected when yield, plant height, spike length, kernels/spike and kernel weight were included in the index.