Fusarium Head Blight of Common Polish Winter Wheat Cultivars – Comparison of Effects of Fusarium avenaceum and Fusarium culmorum on Yield Components

The effects of Fusarium avenaceum and Fusarium culmorum on the reduction in yield components, after independent inoculation of 14 winter wheat cultivars, were investigated. Single isolates of F. avenaceum and F. culmorum were independently used in inoculations of winter wheat heads. Reductions in the following yield traits: 1000‐kernel weight (TKW), the weight (WKH) and number (NKH) of kernels per head after inoculation were analysed statistically. The results indicate differences between both pathogens in their effects on yield traits. The statistical calculations were performed using analysis of variance (a three‐factor experiment) for particular yield trait reductions and multivariate analysis of variance for the yield trait reductions jointly. Almost all of the univariate and multivariate hypotheses concerning no differences between pathogens (F. culmorum, F. avenaceum), climatic conditions (years) and cultivars as well as hypotheses concerning no interactions between factors (pathogens, years, cultivars) were rejected at least at P= 0.05 significance level. The reduction of yield traits indicated individual reactions of the tested winter wheat cultivars to different pathogens. Among the tested traits the highest influence on the rejection of the hypothesis concerning the equivalence of F. avenaceum and F. culmorum was observed for TKW and WKH. The effect of the pathogen on yield reduction was greater for F. avenaceum than for F. culmorum during 1996 and 1997. A comparison of the cultivars indicated that the Begra cultivar showed the highest tolerance to inoculation with both Fusarium pathogens. Moreover, this genotype as well as several others showed lower tolerance to F. avenaceum rather than to F. culmorum, whereas Elena was the only cultivar with the opposite tendency.     

Anther Extrusion and Fusarium Head Blight Resistance in European Wheat

Anther extrusion has been widely discussed as a factor influencing fusarium head blight (FHB) resistance in wheat. This is despite a paucity of quantitative information on its importance, between cultivars, in contrast to that for heading date and plant height. We describe a method applicable to a plant breeding situation at 10 days postanthesis, for assessing the distinct characteristics of anther retention (anthers held within the spikelet) and trapped anthers (partially extruded and trapped between the lemma and palea of the wheat spikelet). FHB resistance was tested in field experiments in 2004 and 2005. In these experiments designed to resemble applications to a plant breeding selection scheme anther retention was significantly correlated with FHB in 2004 ( r  =   0.26; P   <   0.05) and 2005 ( r  =   0.26; P   <   0.05). A higher proportion of anthers retained relating, albeit weakly, with increased FHB susceptibility in European wheat.     

我国小麦赤霉病成灾原因分析及防控策略探讨

小麦是世界上三大粮食作物之一,是全球30亿以上人口的主粮。近年来,由于各种病虫害危害,全球小麦生产和粮食安全受到严重威胁,其中由禾谷镰刀菌引起的小麦赤霉病是小麦生产上重要的病害之一。此外,病菌会产生多种真菌毒素对人畜生命健康构成严重威胁。化学药剂的使用以及抗病品种的种植可以有效地控制小麦赤霉病的发生。但是,由于高产优质抗病品种匮乏、气候变暖等因素影响,小麦赤霉病在我国小麦主产区频繁暴发;同时,赤霉病菌抗药性产生致使化学农药的防控效果大大降低。从气候变化、耕作制度改变、小麦品种抗性及病菌抗药性等方面,分析了赤霉病暴发成灾的主要原因。在此基础上,结合当前赤霉病防控研究进展以及存在的科学问题,探讨该病害持续绿色防控的对策建议,以期为我国小麦赤霉病的防控研究提供参考。     

小麦抗赤霉病利器——他山之石

赤霉病是我国乃至世界小麦(Triticum aestivum)产区的重要病害, 给农业生产和人畜健康造成重大威胁。分离鉴定优质抗病基因、培育抗病品种, 是控制我国麦区赤霉病的重要手段。最近, 山东农业大学孔令让团队完成了二倍体长穗偃麦草(Thinopyrum elongatum)基因组的组装, 并在此基础上通过精细定位和图位克隆分离得到来自长穗偃麦草的抗赤霉病基因Fhb7。他们发现Fhb7编码1个谷胱甘肽转移酶, 对禾谷镰孢菌(Fusarium graminearum)分泌的包括呕吐毒素等在内的多种毒素具有解毒作用, 是1个广谱持久抗病基因。他们还发现Fhb7很可能最初源于内生真菌, 经过基因水平转移进入到偃麦草基因组中。此外, Fhb7不影响其它农艺性状, 且其抗性不受小麦遗传背景影响。这一系列工作揭示了作物抗病演化中的全新机制, 对小麦抗赤霉病育种以及更好地利用长穗偃麦草的丰富基因资源都具有重要意义。     

Differentially Expressed Proteins Associated with Fusarium Head Blight Resistance in Wheat

Background

Fusarium head blight (FHB), mainly caused by Fusarium graminearum, substantially reduces wheat grain yield and quality worldwide. Proteins play important roles in defense against the fungal infection. This study characterized differentially expressed proteins between near-isogenic lines (NILs) contrasting in alleles of Fhb1, a major FHB resistance gene in wheat, to identify proteins underlining FHB resistance of Fhb1.

Methods

The two-dimensional protein profiles were compared between the Fusarium-inoculated spikes of the two NILs collected 72 h after inoculation. The protein profiles of mock- and Fusarium-inoculated Fhb1⁺NIL were also compared to identify pathogen-responsive proteins.

Results

Eight proteins were either induced or upregulated in inoculated Fhb1⁺NIL when compared with mock-inoculated Fhb1⁺NIL; nine proteins were either induced or upregulated in the Fusarium-inoculated Fhb1⁺NIL when compared with Fusarium-inoculated Fhb1⁻NIL. Proteins that were differentially expressed in the Fhb1⁺NIL, not in the Fhb1⁻NIL, after Fusarium inoculation included wheat proteins for defending fungal penetration, photosynthesis, energy metabolism, and detoxification.

Conclusions

Coordinated expression of the identified proteins resulted in FHB resistance in Fhb1⁺NIL. The results provide insight into the pathway of Fhb1-mediated FHB resistance.     

Influence of Growth Stage on Fusarium Head Blight and Deoxynivalenol Production in Wheat

Fusarium head blight is a major concern for wheat production worldwide. The fungi that cause the disease may infect head tissues from flowering to late stages of kernel development, but a better understanding of the influence of the time of infection on grain weight reduction and mycotoxin accumulation resulting from the infection process is needed. We investigated the influence of wheat reproductive stage at the time of inoculation on disease and grain quality parameters, especially production of deoxynivalenol (DON) in mature grains. Heads of Norm wheat were spray inoculated with a macroconidial suspension of a DON‐producing isolate of Fusarium graminearum at each of six reproductive stages from flowering to hard dough. Plants were incubated in a mist chamber for 48 h and then moved to the greenhouse until maturity. Norm wheat was susceptible at all stages inoculated but the highest grain weight reduction and DON accumulation occurred in plants inoculated past flowering to late milk stages. However, high incidences of kernel infection and significant levels of DON accumulation resulted from inoculations as late as the hard dough stage, even though there was no corresponding reduction in grain weight compared to non‐inoculated plants. The occurrence of commercially significant levels of DON in plump, high‐yielding wheat may result from infections that occur during favourable environments well after the flowering stages. Late infection and DON production should therefore be a future research focus for wheat breeding and integrated management of FHB and an important consideration for grading systems that employ the presence of visibly damaged kernels as a means of estimating DON content of wheat.     

Effects of Fusarium graminearum Metabolites on Wheat Tissue in Relation to Fusarium Head Blight Resistance

Fusarium head blight (FHB) of wheat is caused by Fusarium graminearum which produces many secondary metabolites including the trichothecene mycotoxins deoxynivalenol (vomitoxin) and 3-acetyldeoxynivalenol. Coleoptile tissue segements from 14 spring wheat cultivars were exposed to the F. graminearum metabolites deoxynivalenol, 3-acetyldeoxynivalenol, butenolide (all known mycotoxins), sambucinol, culmorin and dihydroxycalonectrin in a bioassay. The tissue of most cultivars was inhibited, at a concentration of 10^?6M by the trichothecenes tested and up to 10^?3M for the other compounds. Deoxynivalenol and 3-acetyldeoxynivalenol, which affect protein synthesis at the ribosome, are therefore potent phytotoxins in addition to being mycotoxins. The resistance or susceptibility of each cultivar to FHB was established in a field experiment. A comparison of the two sets of data indicated that resistant cultivars could tolerate much higher concentrations of the metabolites tested than susceptible cultivars. Some resistant material can tolerate 10 to 1000 times the concentration of the trichothecenes, compared with susceptible cultivars, with no effect on growth. The data suggest that it may be possible to screen germplasm rapidly for FHB resistance in vitro and a new type of resistance in wheat to this disease is proposed based on the apparent insensitivity to trichothecenes by resistant cultivars, additional to the three types of resistance described in the literature.     

小麦抗赤霉病外源种质的创制和育种利用

小麦赤霉病是危害小麦安全生产的重要病害之一,种植抗病品种是防治赤霉病最经济有效的手段。目前在生产上应用的抗源很少,越来越多的研究者将目光转移到小麦的近缘属种,寻找新的抗源以及寻求新的育种突破。携带抗性基因的外源染色体可以通过染色体工程手段以附加系、代换系和易位系等形式导入小麦。综述了将大赖草等多个小麦近缘种的抗赤霉病基因导入普通小麦、创制抗病外源种质和育种利用的最新研究进展,以期为小麦抗赤霉病育种提供参考信息。     

小麦赤霉病新抗源的发掘与抗性位点的检测分析

小麦赤霉病是由禾谷镰刀菌引起的世界性重要病害,发掘优异的抗性种质资源、培育抗病品种是持续防治赤霉病最经济且环境友好的措施。为发掘新的赤霉病抗源,本研究于2017—2021年在弥雾保湿大棚中,采用单花滴注法对642份小麦种质资源的赤霉病抗扩展性进行鉴定,同时利用已知抗赤霉病基因/位点Fhb1~Fhb7的分子标记对筛选出的抗性种质基因型进行分析。结果表明,不同年份间赤霉病病小穗率的相关性均达到极显著水平。筛选到3年及以上赤霉病抗性优于扬麦158的种质81份,主要来自长江中下游麦区,其中33份种质连续4年抗性优于扬麦158;筛选到3年及以上抗性与苏麦3号相当的种质9份,分别为望水白、Grandin、浩麦1号、剑子麦、魁小麦、农林26、软秆洋麦、苏麦2号和武农6号,其中剑子麦、软秆洋麦、苏麦2号和Grandin连续4年抗性与苏麦3号相当。对抗性种质携带的抗赤霉病基因/位点进行分析发现,浩麦1号、冀师7225-28、南农13Y110、石优17和武农6号不携带任何已知抗赤霉病基因/QTL,为小麦抗赤霉病研究和品种培育提供了新的种质资源和理论依据。     

Pathogenic Variation of Fusarium Isolates Associated with Head Blight of Wheat in Australia

This paper examines the level of pathogenic diversity in Australian Fusarium pseudograminearum and Fusarium graminearum isolates for head blight from the assessment of 51 wheat germplasm lines, barley, triticale, rye, maize and sorghum plants. A set of nine putative wheat differentials were selected and assessed with 10 F. graminearum and 12 F. pseudograminearum isolates. Isolates of both species were pathogenic on all the wheat germplasm lines, barley triticale and rye. The isolates differed largely in a quantitative way with only small differential effects and were statistically demarcated into three pathogenicity groups: low, intermediate and high. Such distribution patterns suggest that wheat germplasm lines employ different resistance mechanisms to each group of isolates and the three pathogenicity groups may have different mechanisms controlling pathogenicity. The aggressiveness of F. graminearum and F. pseudograminearum isolates on the wheat germplasm lines were marginally correlated (r = 0.40). Durum wheats were ranked as the most susceptible while Sumai 3, Ituo Komugi, Sotome A, Sotome and Nobeokabouzu komugi were consistently grouped as resistant by both species. These findings reiterate the need to consider pathogen variability in the screening, selection and improvement of resistance to head blight in wheat.     

Minor Differences in Anther Extrusion Affect Resistance to Fusarium Head Blight in Wheat

Fusarium head blight (FHB) remains a serious problem due to yield loss and mycotoxin accumulation in wheat production worldwide. We previously reported that the closed‐flowering (no anther extrusion) characteristic was effective for increasing resistance to FHB infection. In this study, we investigated the relationships between the degree of anther extrusion (AE) and FHB damage using double haploid lines (DHLs), derived from F₁ plants from crosses between closed‐flowering and opened‐flowering varieties. These DHLs exhibited various degrees of AE, and the degree of AE was significantly different among DHLs, regardless of the year and environment (pot‐ or field‐grown). FHB severity was the lowest in closed‐flowering DHLs, and DHLs with partially extruded anthers showed significantly higher FHB symptoms than those with closed‐flowering phenotypes. In general, DHLs with partially extruded anthers also had relatively severe FHB symptoms compared with those exhibiting full anther extrusion. FHB severity was significantly correlated with Fusarium‐damaged kernels and deoxynivalenol concentration. The results of this study showed that partially extruded anthers were considered to be a source of FHB infection. The closed‐flowering phenotype improved resistance to FHB infection. Meanwhile, phenotypes with rapid anther extrusion and ejection also could contribute to the avoidance of FHB infection.     

小麦抗赤霉病品种NBS同源序列克隆与分析

根据二穗短柄草NBS-LRR类基因的保守序列设计同源引物,以小麦抗赤霉病品种苏麦3号、宁7840和望水白基因组DNA为模板,通过PCR扩增,得到43条序列,其中4条为非编码序列或结构域不完整;39条与植物抗病基因同源,其中的7条内部存在终止密码子,可能是假基因,经过比对分析,其余32条具有连续的开放阅读框和保守结构域,推导的氨基酸序列均具有Kinase-1a、Kinase-2和Kinase-3a及GLPL区等几个保守区,在GenBank中均能找到与之高度同源的其他物种的核酸序列,并且Kinase-2的最后一个氨基酸均为色氨酸(W),属于non-TIR类NBS基因。32条序列可分为4大类,它们之间核苷酸同源性为64%-98%,编码氨基酸同源性为22%-98%。根据序列分析随机设计5对不同基因特异性引物,并利用RT-PCR技术进行表达分析,结果表明,7-1、s-3、s-4和w-2均能表达,说明这些片段可能是功能性抗病基因的部分序列;7-13不表达,再次证明属于假基因。32条序列在之前未被报道过,这些RGA可以作为筛选赤霉病功能性抗病基因的候选序列。     

Beauvericin Production by Fusarium Species

Beauvericin is a cyclohexadepsipeptide mycotoxin which has insecticidal properties and which can induce apoptosis in mammalian cells. Beauvericin is produced by some entomo- and phytopathogenic Fusarium species (Fusarium proliferatum, F. semitectum, and F. subglutinans) and occurs naturally on corn and corn-based foods and feeds infected by Fusarium spp. We tested 94 Fusarium isolates belonging to 25 taxa, 21 in 6 of the 12 sections of the Fusarium genus and 4 that have been described recently, for the ability to produce beauvericin. Beauvericin was produced by the following species (with the number of toxigenic strains compared with the number of tested strains given in parentheses): Fusarium acuminatum var. acuminatum (1 of 4), Fusarium acuminatum var. armeniacum (1 of 3), F. anthophilum (1 of 2), F. avenaceum (1 of 6), F. beomiforme (1 of 1), F. dlamini (2 of 2), F. equiseti (2 of 3), F. longipes (1 of 2), F. nygamai (2 of 2), F. oxysporum (4 of 7), F. poae (4 of 4), F. sambucinum (12 of 14), and F. subglutinans (3 of 3). These results indicate that beauvericin is produced by many species in the genus Fusarium and that it may be a contaminant of cereals other than maize.     

Evaluation of Spring and Winter Wheat Reaction to Fusarium culmorum and Fusarium avenaceum

Winter (37), spring (8) wheat accessions and additionally, 7 double haploid (DH) lines were examined for susceptibility to Fusarium seedling blight after inoculation with F. culmorum and F. avenaceum. Winter accessions exhibited lower susceptibility of about 30% to both pathogens than spring cultivars. Susceptibility of winter cultivars varied from low (22%) to high (97%). Evaluation of the root was found to be more reliable than evaluation of coleoptile necrosis.
F. avenaceum infected mostly root and, to a lesser extent, coleoptile and leaves, with about a three times lower disease score of coleoptile against root. F. culmorum caused a 1.5 higher disease score on root than on coleoptile. Susceptibility of DH lines was different from susceptibility of parental forms. Reaction of individual accessions to F. culmorum and F. avenaceum was different.     

Occurrence of beauvericin and enniatins in wheat affected by Fusarium avenaceum head blight.

We evaluated Fusarium contamination and the levels of hexadepsipeptide mycotoxins in 13 wheat samples affected by head blight in Finland. Fusarium avenaceum was the dominant species (91%) isolated from all samples, but isolates of F. culmorum (4%), F. tricinctum (3%), and F. poae (2%) also were recovered. Beauvericin (0.64 to 3.5 microg/g) was detected in all 13 samples. Enniatin B (trace to 4.8 microg/g) was detected in 12 samples, enniatin B(1) (trace to 1.9 microg/g) was detected in 8 samples, and enniatin A(1) (trace to 6.9 microg/g) was detected in 10 samples. Ten of 13 strains of F. avenaceum and 2 strains of F. poae and F. tricinctum produced beauvericin in culture on rice (trace to 70, 9.4, and 33 microg/g, respectively). All strains also produced enniatins (trace to 2,700 microg/g). This is the first report on the natural co-occurence of beauvericin and enniatins in wheat infected predominantly by F. avenaceum.     

The Effect of Nitrogen Fertilization on Fusarium Head Blight Development and Deoxynivalenol Contamination in Wheat

The impact of nitrogen (N) fertilization on the development of Fusarium head blight (FHB) in wheat and the resulting deoxynivalenol (DON) contamination in the kernels was studied. In a first experiment, the disease was assessed on two locations under natural infection pressure. Five different types of nitrogen fertilizer (both organic and mineral) were investigated, each applied at five input rates from 0 to 160 kg N/ha. With all fertilizers, a significant increase of disease intensity was observed with increasing N input, while the type of N fertilizer had poor or no effects on FHB. Depending on the fertilizer used, the percentage of diseased spikelets increased from 2.2% at zero N rate up to 6.6% at 160 kg N input per hectare. In a second series of trials, three spring wheat varieties including one Durum wheat line were artificially inoculated with a Fusarium graminearum and a F. culmorum strain, known producers of DON. A mineral N fertilizer was applied at five input levels from 0 to 160 kg N/ha. A significant increase in FHB intensity and DON contamination in the grain was observed with increasing N from 0 to 80 kg/ha. At higher input rates, relevant in contemporary crop husbandry, disease intensity and toxin contamination remained at constant levels. It is concluded that adaptation of N fertilization represents no relevant tool in managing FHB in practical wheat cultivation.     

Metabolic Biomarker Panels of Response to Fusarium Head Blight Infection in Different Wheat Varieties

Metabolic changes in spikelets of wheat varieties FL62R1, Stettler, Muchmore and Sumai3 following Fusarium graminearum infection were explored using NMR analysis. Extensive 1D and 2D ¹H NMR measurements provided information for detailed metabolite assignment and quantification leading to possible metabolic markers discriminating resistance level in wheat subtypes. In addition, metabolic changes that are observed in all studied varieties as well as wheat variety specific changes have been determined and discussed. A new method for metabolite quantification from NMR data that automatically aligns spectra of standards and samples prior to quantification using multivariate linear regression optimization of spectra of assigned metabolites to samples’ 1D spectra is described and utilized. Fusarium infection-induced metabolic changes in different wheat varieties are discussed in the context of metabolic network and resistance.     

Resistance against Fusarium Head Blight in Transgenic Wheat Plants Expressing the ScNPR1 gene

Fusarium head blight (FHB) is a severe global wheat disease that may cause severe yield losses, especially during epidemic years. Transforming the regulatory genes in the metabolic pathways of disease resistance into wheat via transgenic methods is one way to improve resistance to FHB. ScNPR1 (Secale cereale‐NPR1), a regulatory gene for systemic acquired resistance (SAR), was isolated from S. cereale cv Jingzhouheimai and transformed into the moderately FHB‐susceptible wheat variety Ningmai 13. RT‐PCR analysis indicated that the ScNPR1 gene was stably expressed in transgenic plants. An evaluation of the resistance to FHB revealed that six ScNPR1 transgenic lines (NP1, NP2, NP3, NP4, NP5 and NP6) exhibited significantly higher FHB resistance than the wild‐type wheat Ningmai 13 and the null‐segregated plants. The expression of pathogenesis‐related (PR) genes after Fusarium graminearum inoculation was earlier or higher than those in the wild‐type variety Ningmai 13. The high expression in the early stages of PR genes should account for the enhanced FHB resistance in the transgenic lines. Our results suggest that overexpression of ScNPR1 could be used to improve FHB resistance in wheat.     

sRNA的研究概述及其在小麦赤霉病防治中的应用展望

赤霉病是小麦中最难攻克的真菌性病害之一,不仅能够引起小麦减产,而且会造成DON毒素污染,目前小麦赤霉病的防控仍面临瓶颈。近年来,植物小分子非编码RNA(small noncoding RNA, sRNA)在植物抗病中的作用引起了广泛的关注。sRNA以其分子量小、可移动、靶向更精准等特点,理论上可以作为生物防治赤霉病制剂的主要成分。阐述了sRNA在植物抗病中的研究进展以及利用sRNA特异、高效防治赤霉病的可行性,以期为赤霉病的防治提供新策略。     

Mutual Exclusion between Fungal Species of the Fusarium Head Blight Complex in a Wheat Spike

Fusarium head blight (FHB) is one of the most damaging diseases of wheat. FHB is caused by a species complex that includes two genera of Ascomycetes: Microdochium and Fusarium. Fusarium graminearum, Fusarium culmorum, Fusarium poae, and Microdochium nivale are among the most common FHB species in Europe and were chosen for these experiments. Field studies and surveys show that two or more species often coexist within the same field or grain sample. In this study, we investigated the competitiveness of isolates of different species against isolates of F. graminearum at the scale of a single spike. By performing point inoculations of a single floret, we ensured that each species was able to establish independent infections and competed for spike colonization only. The fungal colonization was assessed in each spike by quantitative PCR. After establishing that the spike colonization was mainly downwards, we compared the relative colonization of each species in coinoculations. Classical analysis of variance suggested a competitive interaction but remained partly inconclusive because of a large between-spike variance. Further data exploration revealed a clear exclusion of one of the competing species and the complete absence of coexistence at the spike level.