小偃麦衍生品系的赤霉病抗性评价

由镰孢属(Fusarium)真菌侵染引起的赤霉病是严重威胁小麦生产的重要病害之一,但小麦育种中可直接利用的抗源非常有限。采用单花滴注法接种赤霉菌株F0609,对来源于中间偃麦草或长穗偃麦草的119份小偃麦衍生品系进行3年6个环境的抗病鉴定,发现平均病小穗率<10%的材料有13份,抗性评价为抗病(R);平均病小穗率介于10%~25%之间的材料有61份,抗性评价为中抗(MR);其余45份材料的平均病小穗率介于25%~50%或>50%,抗性评价为中感或高感(MS和S)。在13份高抗赤霉病材料中,CH16387的抗性显著优于苏麦3号和望水白,CH16371和CH16379的抗性显著优于望水白,其余10个品系与抗病对照苏麦3号和望水白的抗性水平相当。这13份材料分别来自小麦-中间偃麦草部分双二倍体TAI8045和小麦-长穗偃麦草部分双二倍体TAP8430与普通小麦的杂交组合,TAI8045抗性显著优于对照品种望水白, TAP8430与苏麦3号和望水白的抗性相当,而杂交组合中的小麦亲本对赤霉病表现感病,推测这些材料的抗性可能来自TAI8045和TAP8430。这些抗病材料为小麦抗赤霉病育种提供了新的种质资源。     

QTL mapping and marker-assisted selection for Fusarium head blight resistance in wheat: a review

During the past decade, numerous studies have been published on molecular mapping of Fusarium head blight (FHB) resistance in wheat. We summarize the relevant findings from 52 quantitative trait loci (QTL) mapping studies, nine research articles on marker-assisted selection and seven on marker-assisted germplasm evaluation. QTL for FHB resistance were found on all wheat chromosomes except chromosome 7D. Some QTL were found in several independent mapping studies indicating that such QTL are stable and therefore useful in breeding programmes. We summarize and update current knowledge on the genetics of FHB resistance in wheat resulting from QTL mapping investigations and review and suggest FHB breeding strategies based on the available information and DNA markers.     

QTL analysis of resistance to Fusarium head blight in wheat using a 'Wangshuibai'-derived population

Fusarium head blight (FHB) is a devastating disease that reduces the yield, quality and economic value of wheat. For quantitative trait loci (QTL) analysis of resistance to FHB, F₃ plants and F_3:5 lines, derived from a ‘Wangshuibai’ (resistant)/‘Seri82’(susceptible) cross, were spray inoculated during 2001 and 2002, respectively. Artificial inoculation was carried out under field conditions. Of 420 markers, 258 amplified fragment length polymorphism and 39 simple sequence repeat (SSR) markers were mapped and yielded 44 linkage groups covering a total genetic distance of 2554 cM. QTL analysis was based on the constructed linkage map and area under the disease progress curve. The analyses revealed a QTL in the map interval Xgwm533‐Xs18/m12 on chromosome 3BS accounting for up to 17% of the phenotypic variation. In addition, a QTL was detected in the map interval Xgwm539‐Xs15/m24 on chromosome 2DL explaining up to 11% of the phenotypic variation. The QTL alleles originated from ‘Wangshuibai’ and were tagged with SSR markers. Using these SSR markers would facilitate marker‐assisted selection to improve FHB resistance in wheat.     

Types and components of resistance to Fusarium head blight of wheat

Resistance of wheat to Fusarium head blight caused by Fusarium graminearum and F. culmorum was identified in natural epidemics in 1985 and 1987 as well after artificial inoculations (1983–1988 and 1984–1987). Out of 25 genotypes tested, five were identified with no significant difference in head blight scores, but differing significantly in yield after artificial inoculation, i.e. tolerance differences were detected at different resistance levels. Some genotypes that were similar in yield or head blight scores differed in seed infection severity. Genotypes with awns were more susceptible to head blight when tested under natural epidemic condition in the field; but this trait did not influence head blight severity in artificial inoculations. Dwarf genotypes were more severely infected by head blight than tall genotypes under natural conditions, but genotypes of different plant height classes were similarly susceptible after artificial inoculations. In the early generations of a breeding programme resistance measured by visual evaluation of artificial inoculation is the most important way to screen. If selection of dwarf and awned genotypes cannot be avoided, the higher susceptibility caused by awns and dwarfness under natural epidemic conditions can be decreased by a higher level of physiological resistance, as variability in physiological resistance is available. In later generations, traits like percentage of seed infection or tolerance can be identified by additionally measuring yield reduction. Stability of disease reaction appears to be connected with resistance level, the most resistant genotypes are the most stable, and the most susceptible ones tend to have more unstable reactions in different epidemic conditions.     

QTL analysis of resistance to Fusarium head blight in wheat using a 'Frontana'-derived population

Fusarium head blight (FHB or head scab) has become a major limiting factor for sustainable wheat (Triticum aestivum L.) production around the world. For quantitative trait loci (QTL) analysis of resistance to FHB, F₃ plants and F_{3 : 5} lines, derived from a ‘Frontana’ (moderately resistant)/‘Seri82’ (susceptible) cross, were spray‐inoculated in 2001 and 2002, respectively. Artificial inoculations were carried out under field conditions. Of 273 SSR and AFLP markers, 250 could be mapped and they yielded 42 linkage groups, covering a genetic distance of 1931 cM. QTL analysis was based on the constructed linkage map and area under the disease progress curve (AUDPC). The analyses revealed three consistent QTLs associated with FHB resistance on chromosomes 1BL, 3AL and 7AS explaining 7.9%, 7.7% and 7.6% of the phenotypic variation, respectively, above 2 years. The results confirmed the previously described resistance QTL of ‘Frontana’ on chromosome 3AL. A combination of ‘Frontana’ resistance with ‘Sumai‐3’ resistance may lead to lines with augmented resistance expression.     

Nature of wheat resistance to Fusarium head blight and the role of deoxynivalenol for breeding

Twenty (1990-93) and 25 (1994-96) wheat genotypes with different degrees of resistance and origins were tested with seven and eight isolates, respectively, of Fusarium graminearum and four Fusarium culmorum isolates of diverse origin in Europe. Infection severity depended largely on the genotypes and the isolates used. Head blight values, yield response and kernel infection values revealed close but varying relationships with deoxynivalenol (DON) content. This variability is explained by the presence of tolerance mechanisms which affect the relationship between Fusarium head blight severity and yield response. Kernel infection resistance accounted for decreasing Fusarium head blight values. Genotypes were found with lower infection severity and higher DON contamination and vice versa. Evidently, the cultivar has a significant influence on DON production in the infected tissue, i.e. highly susceptible genotypes may have moderate or low accumulation of DON. However, in the most resistant genotypes showing no infection to any of the isolates or only sporadic symptom development, no or very low accumulation of DON was detected. Resistant genotypes gave a stable reaction with b-values close to zero for all traits tested. Susceptible genotypes were unstable under different epidemic conditions and their stability was different for the traits investigated. Therefore, the mean of b-values is suggested to better describe the stability of the wheat genotypes. Significant positive relationships were found between aggressiveness of the isolates and their production of DON in the infected grain. The correlation improved significantly for the nivalenol-producing isolate (F89.4 from France) when the sum of DON and nivalenol contents were considered. This indicates that the total trichothecene toxin-producing capacity of the isolates may be a decisive component of pathogenicity. Since the tests included isolates from different European countries the results provide further proof that no host specificity exists within these pathogens in Europe. This was also valid for kernel infection, yield response and DON accumulation. Therefore, the nature of resistance is horizontal. The results also support the view that there is no difference between the resistance of the host plant to F. graminearum and to F. culmorum.     

Comparison of rye,triticale, durum wheat and bread wheat genotypes for Fusarium head blight resistance and deoxynivalenol contamination

Small-grain winter cereal crops can be infected with Fusarium head blight (FHB) leading to mycotoxin contamination and reduction in grain weight and quality. Although a number of studies have investigated the genetic variation of genotypes within each small-grain cereal, a systematic comparison of the winter crops rye, triticale, durum and bread wheat for their FHB resistance, Fusarium-damaged kernels (FDK) and deoxynivalenol (DON) contamination across species is still missing. We have therefore evaluated twelve genotypes each of four crops widely varying in their FHB resistance under artificial infection with one DON-producing F. culmorum isolate at constant spore concentrations and additionally at crop-specific concentrations in two environments. Rye and triticale were the most resistant crops to FHB followed by bread and durum wheat at constant and crop-specific spore concentrations. On average, rye accumulated the lowest amount of DON (10.08 mg/kg) in the grains, followed by triticale (15.18 mg/kg) and bread wheat (16.59 mg/kg), while durum wheat had the highest amount (30.68 mg/kg). Genotypic variances within crops were significant (p ≤ .001) in most instances. These results underline the differing importance of breeding for FHB resistance in the different crops.     

Expression of a Triticum turgidum var. dicoccoides source of Fusarium head blight resistance transferred to synthetic hexaploid wheat

Yield and quality reductions caused by Fusarium head blight (FHB) have spurred spring wheat (Triticum aestivum L.) breeders to identify and develop new sources of host plant resistance. Four wheat synthetic hexaploids (×Aegilotriticum sp.) were developed, each having a quantitative trait locus (QTL), Qfhs.ndsu‐3AS, providing FHB resistance from Triticum turgidum L. var. dicoccoides chromosome 3A. Synthetics were produced by hybridizing a ‘Langdon’‐T. dicoccoides‐ recombinant chromosome 3A substitution line (2n = 4x = 28, AABB with two accessions of T. tauschii (2n= 2x = 14, DD). Synthetics were inoculated and evaluated for FHB resistance in two separate greenhouse seasons. One synthetic, 01NDSWG‐5, exhibited FHB severity ratings of 36% and 32% in the separate seasons, compared with ratings of 9% and 30% for ‘Alsen’, a FHB‐resistant spring cultivar, and ratings of 70% and 96% for ‘McNeal’, a susceptible spring cultivar, respectively. Synthetic × Alsen backcross‐derived lines were produced to initiate combining different sources of FHB resistance.     

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.     

Quantitative trait loci for resistance to Fusarium head blight in recombinant inbred wheat lines from the cross Huapei 57-2 / Patterson

Fusarium head blight (FHB), primarily caused by Fusarium graminearum in North America, often results in significant losses in yield and grain quality of wheat (Triticum aestivum L.). Evaluation of FHB resistance is laborious and can be affected by environmental conditions. The development of DNA markers associated with FHB quantitative trait loci (QTL) and their use in breeding programs could greatly enhance selection. The objective of this study was to identify the location and effect of QTLs for FHB resistance using simple sequence repeat (SSR) markers. A population of wheat recombinant inbred lines derived from the cross ‘Huapei57-2’/‘Patterson’ was characterized for type II resistance in one field experiment and two tests under controlled conditions in the greenhouse. Bulked segregant analysis followed by QTL mapping was used to identify the major segregating QTLs. Results indicate that ‘Huapei 57-2’ may have the same resistance allele as ‘Sumai3’ at a QTL located on the short arm of chromosome 3B. Other QTLs of lower effect size were identified on the long arm of 3Band on chromosomes 3A and 5B. Our findings along with results from other studies demonstrate that the effect of the QTL on3BS is large and consistent across a wide range of genetic backgrounds and environments. Pyramiding this QTL with other FHB QTLs using marker-assisted selection should be effective in improving FHB resistance in a wheat breeding program. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genotypic and environmental variation of resistance to head blight in triticale inoculated with Fusarium culmorum

Fusarium head blight (FHB) is a widespread disease of small‐grain cereals and can cause substantial losses in grain yield. To assess quantitative genetic parameters as a basis for an efficient breeding programme for resistance, 100 triticale (×Triticosecale Wittm.) genotypes were tested in various environments and artificially inoculated at anthesis with an aggressive isolate of Fusarium culmorum. A visual rating (1–9 scale) was used to assess head blight infection. Five grain yield traits relative to an uninoculated control were also measured. The mean value of the average rating, calculated from four or five readings, was 4.4. It ranged from 3.0 to 5.9 and showed continuous variation. Infection caused a 48% reduction of mean kernel weight per spike, which was the result of 26% fewer kernels per spike and a 32% lower 1000‐kernel weight. The 50‐ml kernel weight was affected by only 20%. The range and genotypic variation was highest for relative kernel weight per spike. For all relative grain yield traits, the most important source of variation was the environment, followed by genotype‐environment interaction, with genotype generally coming last. In contrast, genotypic variation was the most important factor for the disease rating, which also had the highest heritability (h²= 0.89). Phenotypic correlations between the average head blight rating and relative grain yield traits were moderate (r = 0.42–0.57). In conclusion, an average disease rating provides a quantitative assessment of resistance and is suitable for screening large numbers of genotypes. Relative kernel weight per spike gives a ranking of the genotypes that is very similar to the visual score.     

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.     

Molecular mapping of quantitative trait loci for field resistance to Fusarium head blight in a European winter wheat population

Fusarium head blight (FHB), one of the most destructive diseases of wheat in many parts of the world, can reduce the grain quality due to mycotoxin contamination up to rejection for usage as food or feed. Objective of this study was to map quantitative trait loci (QTL) associated with FHB resistance in the winter wheat population ‘G16‐92’ (resistant)/‘Hussar’. In all, 136 recombinant inbred lines were evaluated in field trials in 2001 and 2002 after spray inoculation with a Fusarium culmorum suspension. The area under disease progress curve was calculated based on the visually scored FHB symptoms. For means across all environments two FHB resistance QTL located on chromosomes 1A, and 2BL were identified. The individual QTL explained 9.7% and 14.1% of the phenotypic variance and together 26.7% of the genetic variance. The resistance QTL on 1A coincided with a QTL for plant height in contrast to the resistance QTL on 2BL that appeared to be independently inherited from morphological characteristics like plant height and ear compactness. Therefore, especially the QTL on 2BL could be of great interest for breeding towards FHB resistance.     

Chromosomal location of Fusarium head blight resistance genes and analysis of the relationship between resistance to head blight and brown foot rot

In order to identify chromosomes involved in resistance to Fusarium head blight, a set of 21 substitution lines of Triticum macha (resistant) chromosomes into ‘Hobbit’'sib’(susceptible) were evaluated in trials over 2 years. For the first year's trial, all plants were inoculated on the same day with a conidial suspension of F. culmorum. For the second trial, individual plants were inoculated precisely at mid anthesis of each plant over a period of 2 weeks. The disease level was assessed by visual scoring, relative ear weight and F. culmorumn‐specfic quantitative polymerase chain reaction. The results showed that T. macha chromosomes 1B, 4A and 7A conferred good overall resistance, suggesting that they carry important genes for resistance. In two additional trials, T. macha and ‘Hobbit’'sib’ were evaluated for resistance to brown foot rot. The results showed that T. macha was more susceptible than ‘Hobbit’‘sib’, indicating that stem base disease response is not correlated with head blight resistance in these cultivars.     

Fusarium head blight resistance derived from Lophopyrum elongatum chromosome 7E and its augmentation with Fhb1 in wheat

The objective of this study was to assess the effectiveness of Fusarium head blight (FHB) resistance derived from wheatgrass Lophopyrum elongatum chromosome 7E and to determine whether this resistance can augment resistance in combination with other FHB resistance quantitative trait loci (QTL) or genes in wheat. The ‘Chinese Spring’–Lophopyrum elongatum disomic substitution line 7E(7B) was crossed to three wheat lines: ‘Ning 7840’, L3, and L4. F₂ populations were evaluated for type II resistance with the single‐floret inoculation method in the greenhouse. Simple sequence repeat markers associated with Fhb1 in ‘Ning 7840’ and L4 and markers located on chromosome 7E were genotyped in each population. Marker–trait association was analysed with one‐way or two‐way analysis of variance. The research showed that, in the three populations, the average number of diseased spikelets (NDS) in plants with chromosome 7E is 1.2, 3.1 and 3.2, vs. NDS of 3.3, 7.2 and 9.1 in plants without 7E, a reduction in NDS of 2.1, 4.1 and 5.9 in the respective populations. The QTL on 7E and the Fhb1 gene augment disease resistance when combined. The effect of the QTL on 7E was greater than that on 3BS in this experiment. Data also suggest that the FHB resistance gene derived from L. elongatum is located on the long arm of 7E.     

小麦品种扬麦16赤霉病抗扩展QTL定位及分析

扬麦系列品种赤霉病抗性在世界范围内得到重视,但其抗性遗传机制尚不清楚。扬麦16是近年来大面积推广的抗赤霉病品种,本研究以扬麦16与中麦895杂交构建的174个双单倍体(doublehaploidlines,DH)系为材料,于2017—2019年连续3年对该群体采用单花滴注进行赤霉病抗扩展鉴定。利用660KSNP芯片构建高密度遗传图谱,共检测到6个抗性QTL,分别位于2DL、3BL、4BS、4DS、5BL和6AS染色体上。除4BS位点外,其他5个抗性等位基因均来源于扬麦16。QFhb.yaas-4DS和QFhb.yaas-6AS均在多年被检测到,可解释8.8%～15.0%的表型变异;QFhb.yaas-2DL、QFhb.yaas-3BL仅在1年被检测到,分别解释10.5%和14.7%的表型变异;QFhb.yaas-5BL和来源于中麦895的QFhb.yaas-4BS仅在1年被检测到且效应仅为6.4%和8.3%。QTL效应分析结果表明,相较于单个位点,多个抗性QTL的聚合可显著降低赤霉病严重度。扬麦16抗赤霉病QTL将为揭示扬麦品种抗性遗传机制及开发相应分子标记奠定基础。     

Inheritance of resistance to Fusarium head blight in the wheat lines 'CJ 9306' and 'CJ 9403'

Fusarium head blight (FHB or scab) caused by Fusarium graminearum is a worldwide serious disease in wheat. Exploitation and genetic studies of elite resistance sources can speed up the development of resistant cultivars. To characterize the inheritance of host plant resistance in two new lines, ‘CJ 9306’ and ‘CJ 9403’, developed from a recurrent selection programme in China, six generations P₁, P₂, F₁, F₂, B₁ and B₂ of four crosses and 137 F_{6 : 7} recombinant inbred lines (RILs) from one cross were evaluated in the greenhouse for scab resistance using single‐floret inoculation. The data of area under disease progress curve (AUDPC) in F₂, backcross (BC) and RIL populations exhibited mono‐modal distributions without clear‐cut demarcations and skewing towards resistance. An additive–dominance model was well‐fitted, additive effects played a predominating role, and dominance effects were also significant. Continuous distributions with two major peaks and one minor peak for the number or percentage of scabby spikelets (NSS or PSS) in segregating populations implied the existence of major genes or quantitative trait loci (QTL) for resistance. The estimates of broad‐sense and narrow‐sense heritabilities based on the six‐generation experiment were 56–76% and 26–67% respectively. The estimates of broad‐sense heritabilities based on anova with RILs were 89–90%. These two improved lines with excellent scab resistance and good agronomic traits are of interest for wheat breeding and production.     

Comparison of evaluation methods for selection of resistance to Fusarium head blight in a recurrent selection programme in wheat (Triticum aestivum L.)

The objectives of this study were to compare efficiency of evaluation for resistance to Fusarium head blight (FHB) under two inoculation methods in a recurrent selection programme. Fifty selected homozygous F₅ fertile lines, from each of five cycles (C0, C1, C2, C3 and C4) of recurrent selection, and two control cultivars were evaluated in a split-plot design in 1995 and 1996 under the soil-surface inoculation with Fusarium graminearum-colonized kernels and the single-floret inoculation with ascospore suspension. Comparison of the two inoculation methods using means, ranges, coefficients of variation, heritabilities and correlations among infected spikelet rate (ISR), reaction index (RI) and disease index (DI) indicated that FHB resistance could be evaluated with similar accuracy and precision using either of the two inoculation methods. Regressions of disease scores in the soil-surface inoculation on disease scores in the single-surface inoculation were positive and highly significant, showing a strong relationship between both inoculation methods for FHB resistance. The percentage of lines with similar performance for FHB disease scores in both inoculation methods was high. The soil-surface inoculation and single-floret inoculation appear to be useful techniques for evaluating numerous individuals of segregating population and screening advanced homozygous lines for FHB resistance in a recurrent selection programme in wheat, respectively.     

Molecular and phenotypic characterization of Chinese wheat (Triticum aestivum) cultivars for resistance to Fusarium head blight

To identify excellent cultivars resistant to Fusarium head blight (FHB), 104 wheat cultivars were tested by single-flower inoculation using two prevalent pathogens from 2018 to 2020. Agronomic traits were also investigated. Six FHB-resistance quantitative trait loci (QTL), Fhb1, Fhb2, Fhb4, Fhb5, Fhb7 and Qfhb.crc-2D, have been assessed using previously reported DNA markers. A diagnostic marker has been used for Fhb1, and indicative markers linked to the other QTL were used. Results showed that (i) 12 (11.5%) cultivars were resistant to two pathogens in 3 years; among them, ‘Shengxuan 6’, ‘Wanhongbian 759’, ‘Yunong 903’ and ‘Yunong 901’ had good agronomic traits. (ii) Among cultivars with one resistance QTL, the severities of cultivars carrying Fhb1 and Qfhb.crc-2D were 2.2 and 2.8, respectively, whereas those of cultivars with Fhb2 or Fhb7 were 3.6. Among cultivars with two resistance QTL, the severities of cultivars with Fhb1 + Fhb4, Fhb1 + Fhb7 and Fhb4 + Fhb5 were 2.2, 3.0 and 3.6, respectively. The severity of five cultivars possessing three or four resistance QTL was below 2.5. Fhb1 and Qfhb.crc-2D showed better resistance effects than other resistance QTL.