春化、光周期和矮秆基因在不同国家小麦品种中的分布及其效应

为促进国外种质资源在我国的有效利用,将14个国家的100份代表性小麦品种在国内的8个代表性地点种植,调查抽穗期、成熟期和株高,并以4个春化基因(Vrn-A1、Vrn-B1、Vrn-D1和Vrn-B3)、1个光周期基因(Ppd-D1a)及2个矮秆基因(Rht-B1b和Rht-D1b)的分子标记检测所有品种的基因型。春化基因Vrn-A1a、Vrn-B1、Vrn-D1和vrn-A1+vrn-B1+ vrn-D1的分布频率分别为8.0%、21.0%、21.0%和64.0%;显性等位变异Vrn-A1a、Vrn-B1和Vrn-D1主要存在于来自中国春麦区及意大利、印度、加拿大、墨西哥和澳大利亚的品种中,这些品种一般为春性类型;春化位点均为隐性等位变异或vrn-A1+vrn-D1+Vrn-B1的品种主要分布在中国冬麦区、美国冬麦区、俄罗斯冬麦区,以及英国、法国、德国、罗马尼亚、土耳其和匈牙利,这些地区的小麦均为冬性类型。秋播时,供试品种均能正常抽穗,且携带春化显性变异的材料较隐性类型抽穗早,显性等位变异表现加性效应,4个春化位点均为隐性变异的一些欧美材料因抽穗太晚在杨凌和成都不能正常成熟;而春播时,显性等位变异基因型抽穗的频率高,隐性等位变异基因型基本不能抽穗。光周期不敏感基因Ppd-D1a的分布频率为68.0%,主要分布在中国、法国、罗马尼亚、俄罗斯、墨西哥、澳大利亚和印度,而光周期敏感等位变异Ppd-D1b主要分布在英国、德国、匈牙利和加拿大等中高纬度地区;携带Ppd-D1a的品种较携带Ppd-D1b的品种抽穗早,大多数Ppd-D1a品种在长日照和短日照条件下均能成熟,大部分Ppd-D1b品种在短日照条件下不能成熟。Rht-B1b和Rht-D1b基因的分布频率分别为43.0%和35.0%,其中Rht-B1b主要分布于美国、罗马尼亚、土耳其、意大利、墨西哥和澳大利亚,Rht-D1b主要分布于中国、德国、英国、意大利和印度。一般来说,一个国家的品种携带Rht-B1b或Rht-D1b之一,而这2个基因在高纬度地区分布频率较低。Rht-B1b、Rht-D1b和Ppd-D1a的降秆作用均达显著水平,Rht-B1b和Rht-D1b的加性效应突出。     

小麦Fhb1基因定位、克隆及其在抗赤霉病育种中利用的研究进展

赤霉病已成为小麦第一大病害,严重影响小麦的产量和品质。目前,抗性品种结合化学防治的方法成为当前防治赤霉病的最有效途径。小麦赤霉病抗性受多种基因控制,其中,Fhb1基因抗性效应最大,且抗性稳定,在小麦赤霉病抗性方面具有重要作用。本文综述了赤霉病主效抗性基因Fhb1的定位、分子标记和克隆技术的最新研究进展。利用选择性回交技术和矮败小麦平台将Fhb1基因导入当地小麦中可提升赤霉病抗性,进而结合化学防治方法为我国小麦安全生产提供保障。     

Common resistance of wheat to members of the Fusarium graminearum species complex and F. culmorum

Fusarium head blight (FHB) is the most important disease of wheat in Central Europe. Although common resistance of wheat against several Fusarium species has been proposed recently, no data were available for the recently described species/lineages of F. graminearum and F. culmorum . In this study, twenty wheat genotypes were tested under field conditions by spraying inocula of isolates of eight species of the F. graminearum species complex, and three lineages of F. culmorum in 2003–2004. The severity of FHB, Fusarium damaged kernels, yield reduction and deoxynivalenol/nivalenol contamination were measured. F. culmorum isolates were in general more aggressive to wheat than those belonging to the F. graminearum species complex. The various wheat genotypes exhibited similar reactions against the different Fusarium isolates, indicating that resistance to F. graminearum sensu stricto was similar to that for the other species of the F. graminearum species complex examined. This is an important message to breeders as the resistance relates not only to any particular isolate of F. graminearum , but similarly to isolates of other Fusarium species.     

Selection for Fusarium head blight resistance in early generations reduces the deoxynivalenol (DON) content in grain of winter and spring wheat

Fusarium head blight (FHB) results in yield losses and contamination of kernels by mycotoxins, particularly deoxynivalenol (DON). For minimizing DON content in grain, indirect selection methods would increase gains from selection compared to the costly and time‐consuming DON analysis. The aim of this study was to examine whether an early selection for fewer FHB symptoms would lead to a reduced DON content in grain after inoculation with Fusarium culmorum. Starting with a double‐cross derived population of about 1,100 genotypes, 30 F_1:3 genotypes were selected for FHB rating in a two‐step selection in spring wheat with the non‐adapted resistance sources CM82036 and ‘Frontana’. In winter wheat, 30 F_1:2 genotypes were selected out of a double‐cross derived population of about 600 F₁ plants from crosses with German resistance sources (‘Dream’, G16‐92). Selected genotypes were grouped in three categories according to their FHB rating (low, moderate and high) and analysed afterwards for grain DON content. The three groups differed in their DON content illustrating that indirect selection should already be feasible in the earliest generations. Because of the wide genotypic ranges for DON contents within one grouping, a final DON analysis for selected materials is advisable to achieve full selection gain.     

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.     

山东小麦品种中矮秆基因Rht-B1b、Rht-D1b分布的分子鉴定

利用小麦矮秆基因Rht—B1b、Rht—D1b的4对特异分子标记,NH—BF和MR1、NH—BF和WR1．2、DF和MR2、DF和WR2,对山东小麦品种中矮秆基因Rht—B1b及Rht—D1b的分布情况进行了分子标记鉴定,结果表明在鉴定的150个品种中有20．67％含有Rht—B1b基因,有54．00％含有Rht—D1b基因,二者合计为74．67％;同时含有Rht—B1b和Rht—D1b2个矮秆基因的仅占3．33％。表明山东小麦矮化育种中Rht—D1b基因的利用远高于Rht—B1b。     

Multi-environment evaluation of level and stability of FHB resistance among parental lines and selected offspring derived from several European winter wheat mapping populations

During 2 years and at five locations in Europe, 56 winter wheat genotypes were evaluated for resistance to Fusarium head blight (FHB). The genotypes were both parents and selected recombinants taken from the following populations previously tested for FHB resistance: 'Arina'/'Forno', 'Arina'/'Riband', 'Dream'/'Lynx', G16-92/'Hussar', 'Renan'/'Récital', SVP-72017 × 'Capo' and 'Capo'/'Sumai-3'. In addition, a few control lines were included. FHB resistance was evaluated in replicated experiments under artificial inoculation, disease severity was assessed by repeated visual scorings. The highest level of FHB resistance was found in lines selected from crosses of FHB-resistant winter wheat × 'Sumai-3'. The best lines selected from crosses of moderately resistant winter wheat with susceptible winter wheat were similar in their resistance response to the resistant parent. The level of FHB resistance was correlated with stability of resistance. Susceptible wheat lines tended to exhibit severe symptoms under high disease pressure. The symptoms on resistant lines remained comparatively low even under high disease pressure.     

Confirmation of the relationship between plant height and Fusarium head blight resistance in wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) by QTL meta-analysis

Fusarium head blight (FHB) is a serious wheat disease all over the world. In this study, the relationships between plant height (PH) and FHB were investigated across the whole wheat genome by QTL meta-analysis from fifty-six experiments. Coincident meta-QTL (MQTL) for PH and FHB were found on chromosomes 2D, 3A, 4B, 4D and 7A. Rht-B1, Rht-D1, Rht8, MQTLs P7 and P26 were consistent with FHB MQTLs. The meta-analysis results confirmed the negative associations of Rht-B1, Rht-D1, and Rht8 with FHB resistance. The associations of PH and FHB resistance on chromosomes 3A and 7A have not been reported and need further investigation. These regions should be given attention in breeding programs. MQTLs derived from several resistance sources were also observed. Some FHB MQTLs for different types of resistance overlapped. These findings could be useful for improving wheat varieties with resistance to FHB.     

Effects of genotype and genotype—environment interaction on deoxynivalenol accumulation and resistance to Fusarium head blight in rye, triticale, and wheat

Fusarium culmorum is one of the most important Fusarium species causing head blight infections in wheat, rye, and triticale. It is known as a potent mycotoxin producer with deoxynivalenol (DON), 3‐acetyl deoxynivalenol (3‐ADON), and nivalenol (NIV) being the most prevalent toxins. In this study, the effect of winter cereal species, host genotype, and environment on DON accumulation and Fusarium head blight (FHB) was analysed by inoculating 12 rye, eight wheat, and six triticale genotypes of different resistance levels with a DON‐producing isolate at three locations in 2 years (six environments). Seven resistance traits were assessed, including head blight rating and relative plot yield. In addition, ergosterol, DON and 3‐ADON contents in the grain were determined. A growth‐chamber experiment with an artificially synchronized flowering date was also conducted with a subset of two rye, wheat and triticale genotypes. Although rye genotypes were, on average, affected by Fusarium infections much the same as wheat genotypes, wheat accumulated twice as much DON as rye. Triticale was least affected and the grain contained slightly more DON than rye. In the growth‐chamber experiment, wheat and rye again showed similar head blight ratings, but rye had a somewhat lower relative head weight and a DON content nine times lower than wheat (3.9 vs. 35.3 mg/kg). Triticale was least susceptible with a five times lower DON content than wheat. Significant (P = 0.01) genotypic variation for DON accumulation existed in wheat and rye. The differences between and within cereal species in the field experiments were highly influenced by environment for resistance traits and mycotoxin contents. Nevertheless, mean mycotoxin content of the grain could not be associated with general weather conditions in the individual environments. Strong genotype‐environment interactions were found for all cereal species. This was mainly due to three wheat varieties and one rye genotype being environmentally extremely unstable. The more resistant entries, however, showed a higher environmental stability of FHB resistance and tolerance to DON accumulation. Correlations between resistance traits and DON content were high in wheat (P = 0.01), with the most resistant varieties also accumulating less DON, but with variability in rye. In conclusion, the medium to large genotypic variation in wheat and rye offers good possibilities for reducing DON content in the grains by resistance selection. Large confounding effects caused by the environment will require multiple locations and/or years to evaluate FHB resistance and mycotoxin accumulation.     

小麦–黑麦易位系T1BL·1RS在小麦品种中的分布及其与小麦赤霉病抗性的关联

黑麦1R染色体短臂(1RS)携带条锈病、叶锈病、秆锈病、白粉病和蚜虫等抗性基因。为了检测1RS上是否携带与赤霉病抗性相关的基因,本研究采用1RS特异标记Xscm9对192个来自不同国家的品种/系构成的小麦自然群体和1个重组自交系(RIL)群体(宁7840与Chokwang杂交的F_7群体,共184个系)进行了分子检测,并在2011 2013年采用单花滴注法于温室中进行赤霉病抗性鉴定。结果发现,自然群体中22个品种携带1RS,携带1RS的株系三季赤霉病平均病小穗率(PSS)均显著低于不携带1RS株系的PSS(P0.01),表明1RS对降低病小穗率有显著作用。分子标记和基因组原位杂交(GISH)检测结果表明,宁7840携带1RS。通过对宁7840/Chokwang衍生的RIL群体进行赤霉病抗性鉴定和基因型分析,发现不论主效赤霉病抗性基因Fhb1(标记Xsts142)存在与否,携带1RS株系的PSS显著低于不携带1RS株系的PSS(P0.01);方差分析表明,宁7840携带的Fhb1与1RS在赤霉病抗扩展性上无显著互作(P0.05)。因此认为,黑麦1RS染色体很可能携带赤霉病扩展抗性相关基因,与Fhb1基因有累加效应。     

The potential of genomic‐assisted breeding to improve Fusarium head blight resistance in winter durum wheat

Durum wheat is the most important tetraploid wheat mainly used for semolina and pasta production, but is notorious for its high susceptibility to Fusarium head blight (FHB). Our objectives were to identify and characterize quantitative trait loci (QTL) in winter durum and to evaluate the potential of genomic approaches for the improvement of FHB resistance. Here, we employed an international panel of 170 winter and 14 spring durum lines, phenotyped for Fusarium culmorum resistance at five environments. Heading date, plant height and mean FHB severity showed significant genotypic variation with high heritabilities and FHB resistance was negatively correlated with both heading date and plant height. The dwarfing gene Rht‐B1 significantly affected FHB resistance and the genome‐wide association scan identified eight additional QTL affecting FHB resistance, explaining between 1% and 14% of the genotypic variation. A genome‐wide prediction approach yielded only a slightly improved predictive ability compared to marker‐assisted selection based on the four strongest QTL. In conclusion, FHB resistance in durum wheat is a highly quantitative trait and in breeding programmes may best be tackled by classical high‐throughput recurrent phenotypic selection that can be assisted by genomic prediction if marker profiles are available.     

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.     

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.     

Utilization of alien genes to enhance Fusarium head blight resistance in wheat – A review

Fusarium head blight (FHB) is a destructive disease of wheat worldwide. Sources of resistance to FHB are limited in wheat. Search for novel sources of effective resistance to this disease has been an urgent need in wheat breeding. Fusarium head blight resistance has been identified in relatives of wheat. Alien chromatin carrying FHB resistance genes has been incorporated into wheat through chromosome addition, substitution, and translocation. Relatives of wheat demonstrate a great potential to enhance resistance of wheat to FHB.     

Use of non‐adapted quantitative trait loci for increasing Fusarium head blight resistance for breeding semi‐dwarf wheat

Semi‐dwarf wheat is an important prerequisite for releasing a successful commercial cultivar in high‐yielding environments. In Northern Europe, this aim is achieved by using one of the dwarfing genes Rht‐B1 (formerly known as Rht‐1) or Rht‐D1 (Rht‐2). Both genes, however, result in a higher susceptibility to Fusarium head blight (FHB). We analysed the possibility to use the two non‐adapted FHB resistance quantitative trait loci Fhb1 and Fhb5 (syn. QFhs.ifa‐5A) to counterbalance the negative effect of the dwarfing allele Rht‐D1b in a winter wheat population of 585 doubled‐haploid (DH) lines segregating for the three loci. All lines were inoculated with Fusarium culmorum at four locations and analysed for FHB severity, plant height, and heading date. The DH population showed a significant (p < 0.001) genotypic variation for FHB severity ranging from 3.6% to 65.9% with a very high entry‐mean heritability of 0.95. The dwarfing allele Rht‐D1b reduced plant height by 24 cm, but nearly doubled the FHB susceptibility (24.74% vs. 12.74%). The resistance alleles of Fhb1 and Fhb5 reduced FHB susceptibility by 6.5 and 11.3 percentage points, respectively. Taken all three loci together, Fhb5 alone was already able to reduce FHB susceptibility to the same extent as Rht‐D1b increased it. This opens new avenues for selecting semi‐dwarf wheat by marker‐assisted introgression of Fhb5 without the enhancement of FHB susceptibility.     

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.     

Combined inoculation of wheat pathogens Zymoseptoria tritici and Fusarium culmorum as a tool for increasing selection intensity in resistance breeding

Resistances to Septoria tritici blotch (STB) and Fusarium head blight (FHB) are important goals in European wheat breeding. We tested 25 winter wheat cultivars differing in their resistance to both diseases by inoculating Zymoseptoria tritici or Fusarium culmorum either separately on different plots or combined on the same plot. Experiments were carried out across three location × year combinations in four variants: non‐inoculated, STB inoculated, FHB inoculated and STB+FHB inoculated at the respective optimal plant stages. On the individually inoculated plots, mean STB severities ranged from 12% to 70% and mean FHB severities from 0.3% to 67% across wheat cultivars. The resistances to STB and FHB were not correlated. Mean disease severities of the respective inoculation variants, STB vs. STB+FHB and FHB vs. STB+FHB, were not significantly different (P > 0.1), and correlations between both inoculation variants were extremely high (r = 0.98) for STB. In conclusion, breeding populations have to be selected for both resistances separately, but phenotyping can be performed on the same plot without ranking differences of the respective resistance.     

小麦近缘野生植物的赤霉病抗源筛选及其利用

赤霉病是禾谷镰刀菌等真菌侵染所造成的生育后期的气候型病害。世界各产麦区均有此病害发生,在气候潮湿、温暖多雨的地区尤为严重。培育抗病品种是解决赤霉病危害的根本途径。在披碱草、偃麦草、山羊草和鹅冠草等小麦近缘种属植物中已经鉴定出赤霉病抗性基因。携带赤霉病抗性基因的外源染色体可以通过附加、代换和易位导入小麦。本文综述了小麦近缘野生植物的赤霉病抗源,以及利用这些抗源进行小麦赤霉病抗性育种的研究进展。     

Combining ability of resistance to head blight caused by Fusarium culmorum (W.G. Smith) in the F1 of a seven parent diallel of winter wheat (Triticum aestivum L.)

Fusarium head blight (FHB, scab) caused by Fusarium spp. is a widespread disease of cereals causing relevant yield and quality losses and contaminating cereal products with mycotoxins. Breeding resistant cultivars is the method of choice for controlling the disease. Resistance to FHB is a quantitative trait and is most likely governed by several genes. We present the results of an F₁ diallel analysis of FHB resistance involving six resistant and one susceptible European winter wheat genotypes of diverse origin in order to identify promising combinations for the selection of improved cultivars. Parents and F₁s including reciprocals were evaluated for FHB resistance in an artificially inoculated field trial. Two traits were assessed: visual disease symptoms on the heads and the percentage of Fusarium damaged kernels in a harvested sample. General combining ability (GCA) and specific combining ability (SCA) effects were statistically significant for visual symptoms and kernel damage, whereas reciprocal effects were small or not significant. Heterosis for resistance was common, indicating that the parental genotypes possess different resistance genes. Selection of transgressive segregates should be feasible from such heterotic combinations. This revised version was published online in August 2006 with corrections to the Cover Date.     

Phenotypic selection for high resistance to Fusarium head blight after introgression of quantitative trait loci (QTL) from exotic spring wheat and verification by simple sequence repeat markers a posteriori

Fusarium head blight (FHB) has become an important disease of wheat. We introgressed three resistance quantitative trait loci (QTL) alleles on chromosomes 3B, 5A (from CM82036) and 3A (from ‘Frontana’) into European elite spring wheat and performed phenotypic selection among double‐cross (DC) derived progeny in generations DCF₂ and DCF₃. After recombination and selfing, we analysed 135 phenotypically selected progeny by simple sequence repeat (SSR) markers linked to the QTL. In a second experiment, we forwarded the best 20 progeny for a further two generations by pedigree selection. Progeny were inoculated at two to four locations with Fusarium culmorum and the percentage of infected spikelets per plot was estimated. Both experiments show that phenotypic selection was highly effective. One‐hundred out of 135 phenotypically selected DCF_1:3 progeny had the combination of donor‐QTL alleles (3B + 5A + 3A, 3B + 5A) with the highest effects on FHB resistance. In the subsequent generations, sufficient genotypic variance was detected. The best F_5:7 bulks had similar resistance to the donor CM82036. The FHB rating was reduced in total by 45% points compared to the parental mean. QTL with high effects can be detected solely by phenotypic selection after targeted introgression.