Identification of eight Berberis species from the Yunnan-Guizhou plateau as aecial hosts for Puccinia striiformis f.sp.tritici,the wheat stripe rust pathogen

Puccinia striiformis Westend.f.sp.tritici Erikss.(Pst)infects wheat and causes stripe rust.The rust is heteroecious with wheat as the primary uredinial and telial host and barberry(Berberis spp.)as the alternate pycnial and aecial host.More than 40 Berberis species have been identified as alternate hosts for Pst,and most of these are Chinese Berberis species.However,little is known about Berberis species or their geographic distributions in the Yunnan-Guizhou plateau in southwestern China.The Yunnan-Guizhou plateau is considered to be an important and relatively independent region for the evolution of the wheat stripe rust pathogen in China because the entire disease cycle can be completed within the region.In this study,we conducted a survey of barberry plants in the Yunnan-Guizhou plateau and identified the eight Pst-susceptible Berberis species under controlled conditions,including B.julianae,B.tsienii,B.veitchii,B.wilsonae,B.wilsonae var.guhtzunica,B.franchetiana,B.lepidifolia and B.pruinosa.These species are reported here for the first time to serve as alternate hosts for the wheat stripe rust pathogen under controlled conditions.

Association Analysis of SP-SNPs and Avirulence Genes in Puccinia striiformis f. sp. tritici, the Wheat Stripe Rust Pathogen

Puccinia striiformis f. sp. tritici (Pst) is one of the pathogenic fungi on wheat, caused stripe rust that is a great threat for wheat production all over the world. Intensive efforts have been made to study genetics of wheat resistance to this disease, but few on avirulence of the pathogen due mainly to the nature of obligate biotrophism and the lack of systems for studying its genetics and molecular manipulations. To overcome these limitations, a natural Pst population comprising 352 isolates representative of a diverse virulence spectrum was genotyped using 97 secreted protein-single nucleotide polymorphism (SP-SNP) markers to identify candidate avirulence genes using association analysis. Among avirulence genes corresponding to 19 resistance genes, significantly associated SP-SNP markers were detected for avirulence genes AvYr1, AvYr2, AvYr6, AvYr7, AvYr8, AvYr44, AvYrExp2, AvYrSP, and AvYrTye. These results indicate that association analysis can be used to identify markers for avirulence genes. This study has laid the foundation for developing more SP-SNPs for mapping avirulence genes using segregating populations that can be generated through sexual reproduction on alternate hosts of the pathogen.

Variation in Telial Formation of <i>Puccinia striiformis</i>in the United States

Puccinia striiformis, the causal agent of stripe rust in wheat, barley, and various wild grasses, produces urediniospores and teliospores on these primary or auxiliary hosts. Telial formation, which stops producing infectious urediniospores, is affected by various host and environmental conditions. However, it is not clear if variation exists among different isolates in the United States. To determine the differences in the pathogen population, 1423 isolates collected in the United States from 2013 to 2016 were tested for telial formation in the seedling stage at a diurnal temperature profile of 4&deg;C - 20&deg;C. The percentages of telial formation varied greatly among the isolates. Of the 1423 isolates, 62.97% produced telia with the percentages of telial formation ranging from 1% to 95% under the test conditions. The formation of telia was significantly affected by the year and the month when the isolates were collected. The epidemiological regions or states, host plants (wheat, barley, and grasses), and races of the isolates did not significantly affect telial formation. However, significant effects on telial formation were observed by interactions between year and region, year and race, month and region and among year, month, and region, as well as between year and month. The results showed that telial formation is a complex trait under the genetic control of the pathogen isolates for adaptation to different environments. Further studies are needed to identify genes involved in the formation of telia and the relationship of telial formation to the survival, aggressiveness, fitness, and evolution of the pathogen.

The TaFIM1 gene mediates wheat resistance against Puccinia striiformis f. sp. tritici and responds to abiotic stress

Fimbrin, a regulator of actin cytoskeletal dynamics that participates in numerous physiological and biochemical processes, controls multiple developmental processes in a variety of tissues and cell types. However, the role of fimbrin in pathogen defense of wheat and the mechanisms have not been well studied. Here, we investigated that the expression of TaFIM1 gene of wheat was significantly induced in response to avirulent race of Puccinia striiformis f. sp. tritici(Pst) and silencing of TaFIM1 by virus-induced gene silencing method. The results show that silencing of TaFIM1 resulted in a reduction of resistance against the stripe rust indicated by both phenotypes and a histological examination of Pst growth. Additionally, the expression level of Ta FIM1 gene was up-regulated under abiotic stresses. These findings suggest that Ta FIM1 functions as a positive regulator of pathogen resistance of wheat plants and response to abiotic stress. Our work may show new light on understanding the roles of fimbrin in wheat.

Effects of Different Cultivation Patterns of Wheat on Population Structure of Puccinia striiformis West.f. sp. tritici

The paper was to study the effects of different cultivation patterns( mix cultivation and monocultivation) of wheat on population structure of Puccinia striiformis West. f. sp. tritici in the fields. Five race-specific-markers( CY32,CY31,CY29,CY23 and Shuiyuan pathotype) were used to survey 113 infected samples collected from two cultivation patterns. The results indicated that frequency of race-specific-markers under monocultivation was higher than that under mix cultivation; the dominant race-specific-markers were CY32 and CY29 under monocultivation,and the frequency of detection were 81. 5% and 78. 5%,respectively. The dominant race-specific-markers were CY29 and Shuiyuan pathotype under mix cultivation,and the frequency of detection are 41. 7% and 18. 8%,respectively.Several race-specific-markers were detected in single infected leaf,and 41. 7% of infected single leaf were detected with more than two race-specific-markers,58. 3% of infected single leaf were detected with one race-specific-marker under mix cultivation pattern,while there were 75. 0% infected leaves with more than two race-specific-markers and 25. 0% infected single leaf detected with one race-specific-marker under monocultivation pattern. The results indicated that mix cultivation pattern of wheat can reduce races on single leaf,affect the distribution of races in infected leaves,and suppress the occurrence frequency of dominant races of P. striiformis in the fields significantly,subsequently reduced severity and prevalence of the disease.

The Puccinia striiformis effector Hasp98 facilitates pathogenicity by blocking the kinase activity of wheat TaMAPK4

The obligate biotrophic fungus Puccinia striiformis f.sp.tritici(Pst)employs virulence effectors to disturb host immunity and causes devastating stripe rust disease.However,our understanding of how Pst effectors regulate host defense responses remains limited.In this study,we determined that the Pst effector Hasp98,which is highly expressed in Pst haustoria,inhibits plant immune responses triggered by flg22or nonpathogenic bacteria.Overexpression of Hasp98 in wheat(Triticum aestivum)suppressed avirulent Pst-triggered immunity,leading to decreased H2O2accumulation and promoting P.striiformis infection,whereas stable silencing of Hasp98 impaired P.striiformis pathogenicity.Hasp98 interacts with the wheat mitogenactivated protein kinase TaMAPK4,a positive regulator of plant resistance to stripe rust.The conserved TEY motif of TaMAPK4 is important for its kinase activity,which is required for the resistance function.We demonstrate that Hasp98inhibits the kinase activity of TaMAPK4 and that the stable silencing of TaMAPK4 compromises wheat resistance against P.striiformis.These results suggest that Hasp98 acts as a virulence effector to interfere with the MAPK signaling pathway in wheat,thereby promoting P.striiformis infection.

New insights in the battle between wheat and Puccinia striiformis

Wheat stripe rust caused by Puccinia striiformis f.sp.tritici(Pst)poses a great threat to wheat production worldwide.The rapid change in virulence of Pst leads to a loss of resistance in currently resistant wheat cultivars,which results in frequent disease epidemics.Therefore,a major focus is currently placed on investigating the molecular mechanisms underlying this rapid variation of pathogenicity and coevolving wheat resistance.Limited by the lack of a system for stable transformation of Pst and the difficulties in wheat transformation,it is not easy to generate deeper insights into the wheat-Pst interaction using established genetic methods.Nevertheless,considerable effort has been made to unravel the wheat-Pst interaction and significant progress is being made.Histology and cytology have revealed basic details of infection strategies and defense responses during wheat-Pst interactions,identified cellular components involved in wheat-Pst interactions,and have helped to elucidate their role in the infection process or in plant defense responses.Transcriptome and genome sequencing has revealed the molecular features and dynamics of the wheat-Pst pathosystem.Extensive molecular analyses have led to the identification of major components in the wheat resistance response and in Pst virulence.Studies of wheat-Pst interactions have now entered a new phase in which cellular and molecular approaches are being used.This review focuses on the cellular biology of wheat-Pst interactions and integrates the emerging data from molecular analyses with the histocytological observations.

A secreted catalase contributes to Puccinia striiformis resistance to host-derived oxidative stress

Plants can produce reactive oxygen species(ROS)to counteract pathogen invasion,and pathogens have also evolved corresponding ROS scavenging strategies to promote infection and pathogenicity.Catalases(CATs)have been found to play pivotal roles in detoxifying H_(2)O_(2)formed by superoxide anion catalyzed by superoxide dismutases(SODs).However,few studies have addressed H_(2)O_(2)removing during rust fungi infection of wheat.In this study,we cloned a CAT gene PsCAT1 from Puccinia striiformis f.sp.tritici(Pst),which encodes a monofunctional heme-containing catalase.PsCAT1 exhibited a high degree of tolerance to pH and temperature,and forms high homopolymers.Heterologous complementation assays in Saccharomyces cerevisiae reveal that the signal peptide of PsCAT1 is functional.Overexpression of PsCAT1 enhanced S.cerevisiae resistance to H_(2)O_(2).Transient expression of PsCAT1 in Nicotiana benthamiana suppressed Bax-induced cell death.Knockdown of PsCAT1 using a host-induced gene silencing(HIGS)system led to the reduced virulence of Pst,which was correlated to H_(2)O_(2)accumulation in HIGS plants.These results indicate that PsCAT1 acts as an important pathogenicity factor that facilitates Pst infection by scavenging host-derived H_(2)O_(2).

Determination of heterozygosity for avirulence/virulence loci through sexual hybridization of Puccinia striiformis f. sp. tritici

Wheat stripe rust caused by Puccinia striiformis f. sp. tritici is one of the most devastating diseases of wheat worldwide and resistant cultivars are vital for its management. Therefore, investigating the heterozygosity of the pathogen is important because of rapid virulence changes in isolates heterozygous for avirulence/virulence.An isolate of P. striiformis f. sp. tritici was selfed on Berberis shensiana to determine the heterozygosity for avirulence/virulence loci. One hundred and twenty progeny isolates obtained from this selfing were phenotyped using 25 lines of wheat containing Yrgenes and genotyped with 96 simple sequencing repeat markers, with51 pathotypes and 55 multi-locus genotypes being identified. All of these were avirulent on lines with Yr5,Yr10, Yr15, Yr24 and Yr26 and virulent on lines with Yr17,Yr25 and YrA, indicating that the parental isolate was homozygously avirulent or homozygously virulent for these loci. Segregation was found for wheat lines with Yr1,Yr2, Yr4, Yr6, Yr7, Yr8, Yr9, Yr27, Yr28, Yr32, Yr43, Yr44,YrExp2, YrSp, YrTr1, YrTye and YrV23. The 17 cultivars to which the Pst was identified as heterozygous with respect to virulence/avirulence should not be given priority in breeding programs to obtain new resistant cultivars.

Comments on “Determination of heterozygosity for avirulence/virulence loci through sexual hybridization of Puccinia striiformis f. sp. tritici”

Over the past decades Puccinia striiformis f.sp.tritici（Pst）has developed into one of the most,if not the most important fungal pathogen in wheat production worldwide.In China,Pst has caused numerous epidemics with partially devastating yield losses[1].The occurrence of the'warrior'race in Europe in 2011 also caused significant problems[2].Pst,like other obligate biotrophs,is characterized by a high degree of genetic variability,especially with respect

A small knottin-like peptide negatively regulates in wheat to stripe rust resistance during early infection of wheat

Although Blufensins(Bln)have important functions in the response of plants to biotic stress the precise functioning of Bln in wheat remains largely unknown.Here we isolated a Bln gene(TaBln4)from Suwon 11 infected by Puccinia striiformis f.sp.tritici(Pst).Expression of TaBln4 increased in host plants at the early stage of infection with a virulent Pst race(CYR31)but was unchanged in response to infection by an avirulent race(CYR23).Transcription levels of TaBln4 were also regulated by hormone and abiotic stresses.Expression of TaBln4 in tobacco leaves suppressed Bax-induced programmed cell death.Knockdown of TaBln4 by virus-induced gene silencing inhibited colonization of race CYR31 by increasing the accumulation of H2O2 and formation of hypersensitive responses(HR).Transient overexpression of TaBln4 by a transient overexpression system(BSMV-VOX)increased the susceptibility of wheat to CYR31.Results from bimolecular fluorescence complementation and pull-down assays demonstrated that TaBLN4 interacted with calmodulin.Taken together,our results suggest that TaBln4 negatively regulates resistance in wheat to Pst in a reactive oxygen species(ROS)-and HR-dependent manner.

2017年我国小麦条锈病流行特点及重发原因分析

2017年小麦条锈病在我国黄淮海麦区大范围流行,表现出汉水流域及黄淮南部见病时间早、扩散速度快、黄淮海麦区流行范围广等特点。本文在系统总结2017年全国小麦条锈病流行特点的基础上,分析认为极端暖冬气候、春季多雨适温气候条件和主产麦区缺乏抗性品种等因素是导致2017年我国小麦条锈病大流行的主要原因。

十堰麦区小麦条锈病精准监测及药剂防治的初步研究

通过收集整理历史资料、实地普查、定点监测站和孢子捕捉仪收集条锈菌孢子等方法进行精准监测,初步研究了十堰麦区小麦条锈病(Puccinia striiformis West)冬繁区的核心地带和传播路径;通过大田药剂喷雾和盆栽药剂拌种试验,筛选出适合秦巴山十堰麦区防治小麦条锈病的杀菌剂和剂型,总结出一套适合该麦区小麦条锈病的综合治理技术。

Review Article: High-Temperature Adult-Plant Resistance, Key for Sustainable Control of Stripe Rust

High-temperature adult-plant (HTAP) resistance expresses when plants grow old and the weather becomes warm. This non-race specific and durable type of resistance has been used successfully in control of wheat stripe rust in the US since early 1960s. This article describes practical procedures for identification and characterization of HTAP resistance and reviews recent studies on discovery of genes conferring HTAP resistance. Recent studies providing insights to the molecular basis for the durability of HTAP resistance will be presented. Strategies for improving levels of HTAP resistance and improving control of stripe rust through combining HTAP resistance with effective all-stage resistance will be discussed.

Molecular mapping of stripe rust resistance gene YrH9017 in wheat-Psathyrostachys huashanica introgression line H9017-14-16-5-3

Several new stripe rust pathogen races emerged in the wheat growing regions of China in recent years.These races were virulent to most of the designated wheat seedling resistance genes.Thus,it is necessary and worthwhile to identify new valuable resistant materials for the sake of diversifying resistant sources,pyramiding different resistance genes and achieving durable resistance.Here,we identified the resistance gene,temporarily designated as YrH9017,in wheat-Psathyrostachys huashanica introgression line H9017-14-16-5-3.A total of 146 F2 plants and their derived F2:3 families in a cross of Mingxian 169 and H9017-14-16-5-3 were used to evaluate seedling stripe rust response and as a mapping population.Finally,we constructed a genetic map including eight simple sequence repeat(SSR) markers and expressed sequence tag(EST) markers.YrH9017 was located on the long arm of chromosome 2A and closely linked with two EST-sequence tagged site(EST-STS) markers BG604577 and BE471201 at 1.3 and 1.8 cM distance,respectively.The two closest markers could be used for marker-assisted selection of YrH9017 in breeding.

Effect of stripe rust on the yield response of wheat to nitrogen

Nitrogen(N) is the most important fertiliser element determining the productivity of wheat.N nutrition is known to affect the level of stripe rust infection, with higher N associated with increased disease severity. Stripe rust, caused by Puccinia striiformis f. sp. tritici, is a major yield-limiting disease of wheat in Australia. This paper describes experiments designed to investigate the agronomic response to the interaction of various levels of N application and stripe rust severity in wheat varieties differing in response. Experimental plots were established in crop seasons 2006 and 2007 on the Liverpool Plains of northern NSW, Australia.Yield, biomass, grain protein content(GPC) and harvest index(HI) data were recorded.Increased rates of N increased the severity of stripe rust during grain filling. N application also increased yield and GPC in all varieties in both years. Stripe rust reduced the yield of the rust-susceptible wheat varieties, and GPC and proportion of added N recovered in the grain were also reduced in one year but not the other. It was evident from our experiment that stripe rust caused yield loss accompanied by either no change or reduction in GPC, indicating that the total amount of N entering the grain was reduced by stripe rust. The effects of stripe rust on N yield are most likely associated with reduced uptake of N during grain filling.

Identification of a new stripe rust resistance gene in Chinese winter wheat Zhongmai 175

Stripe rust is a serious foliar disease posing a grave threat to wheat production worldwide. The most economical and environmentally friendly way to control this disease is to breed and deploy resistant cultivars. Zhongmai 175 is an elite winter wheat cultivar conferring resistance to a broad spectrum of Puccinia striiformis f. sp. tritici(Pst) races. To identify the resistance gene in the cultivar, genetic analysis was conducted using the parents, F1, F2 and F2:3 populations derived from the cross of Lunxuan 987/Zhongmai 175. Segregations in the F2 and F2:3 populations indicated a single dominant gene conferring resistance to stripe rust in Zhongmai 175, temporarily designated Yr ZM175. Bulked segregant analysis(BSA) with wheat i Select 90 K SNP array determined a preliminary location of Yr ZM175. Subsequently, Yr ZM175 was mapped on chromosome 2AS using simple sequence repeats(SSR), expressed sequence tags(EST) and newly-developed kompetitive allele specific PCR(KASP) markers, being flanked by Xgwm636 and Xwmc382 at genetic distances of 4.9 and 8.1 c M, respectively. Comparison of reaction patterns of Yr ZM175 on 23 Pst races or isolates and pedigree analysis with other genes on chromosome 2AS suggested that it is likely to be a new gene for resistance to stripe rust. The resistance gene and linked molecular markers will be useful in wheat breeding targeting for the improvement of stripe rust resistance.

Three co-located resistance genes confer resistance to leaf rust and stripe rust in wheat variety Borlaug 100

Leaf rust(LR) and stripe rust(YR) are important diseases in wheat producing areas worldwide and cause severe yield losses under favorable environmental conditions when susceptible varieties are grown. We determined the genetic basis of resistance to LR and YR in variety Borlaug 100 by developing and phenotyping a population of 198 F6 recombinant inbred lines derived from a cross with the susceptible parent Apav#1. LR and YR phenotyping were conducted for 4 and 3 seasons, respectively, at CIMMYT research stations in Mexico under artificial epidemics. Mendelian segregation analyses indicated that 3–5 LR and 2 YR genes conferred resistance in Borlaug 100. Lr46/Yr29(1 BL), Yr17(2 AS) and Yr30(3 BS) were present in the resistant parent and segregated in the RIL population based on characterization by molecular markers linked to these genes. When present alone, Lr46/Yr29 caused average 13% and 16% reductions in LR and YR severities, respectively, in RILs. Similarly, Yr17 and Yr30 reduced YR severities by 57% and 11%, respectively. The Yr30 and the Yr17 translocation were also associated with 27% and 14% reductions, respectively, in LR severity, indicating that the 3 BS and 2 AS chromosomal regions likely carry new slow rusting LR resistance genes, temporarily designated as Lr B1 and Lr B2, respectively. Additive effects of Yr30*Yr17, Yr29*Yr17 and Yr29*Yr30 on YR and LR were significant and reduced YR severities by 56%,55%, and 45%, respectively, and LR severities by 34%, 40%, and 45%, respectively. Furthermore, interaction between the three genes was also significant, with mean reductions of 70% for YR and 54% for LR severities. Borlaug 100, or any one of the 21 lines with variable agronomic traits but carrying all three colocated resistance loci, can be used as resistance sources in wheat breeding programs.

SNP-based linkage mapping for validation of adult plant stripe rust resistance QTL in common wheat cultivar Chakwal 86

Wheat crops in China are constantly challenged by stripe rust. Deployment of cultivars with diverse resistances is the best strategy to control the disease. A recombinant inbred line(RIL) population derived from a cross between the resistant cultivar Chakwal 86 and susceptible landrace Mingxian 169 was studied in multiple environments to examine the underlying genetics and to identify quantitative trait loci(QTL) for stripe rust resistance.One hundred and twenty-eight RILs were genotyped with wheat 35 K SNP array and a genome-wide linkage map with 1480 polymorphic SNP markers, or bins, was constructed.Two major QTL on chromosomes 1BL and 3BS, and one minor QTL on 6BS had significant effects in reducing stripe rust severity. The QTL were validated using composite interval mapping(CIM) and inclusive composite interval mapping(ICIM). These methods explained59.0%–74.1% of the phenotype variation in disease response. The QTL on chromosome 1 BL was confirmed to be Yr29/Lr46 and the one on 3BS was the resistance allele identified in CIMMYT germplasm but was not Yr30/Sr2. The QTL on 6BS probably corresponded to previously known QTL. RILs with combined QTL were more resistant than those with single or no QTL. Kompetitive allele-specific PCR(KASP) assays for the QTL with largest effect QTL on chromosome 3BS were performed on a subset of RILs and 150 unrelated wheat lines. The QTL on 3BS with its linked KASP markers can be used in marker-assisted selection to improve stripe rust resistance in breeding programs.

Effects of UV-B radiation intensity and timing on epidemiological components of wheat stripe rust

Stripe rust caused by Puccinia striiformis f. sp. tritici is an important wheat disease worldwide that is greatly influenced by environmental conditions. Ultraviolet B(UV-B) radiation is one important environmental factor affecting the occurrence and epidemiology of wheat stripe rust. Investigating UV-B radiation effects on the epidemiology of stripe rust may be conducive to monitoring and predicting this disease. In this study, wheat seedlings were exposed to UV-B radiation during different periods under laboratory conditions and radiation effects on epidemiological components of wheat stripe rust were investigated. Results showed that incubation period was shortened, and the infection efficiency, sporulation quantity and disease index increased when UV-B radiation was performed only pre-inoculation. When the UV-B radiation was performed only postinoculation or both pre-and post-inoculation, the incubation period was prolonged, and the infection efficiency, sporulation quantity and disease index were reduced. When healthy wheat seedlings were inoculated using urediospores collected from wheat leaves irradiated by UV-B only post-inoculation or both pre-and post-inoculation, infection efficiency, sporulation quantity and disease index were also reduced. However, in the latter, the disease incubation period did not differ under varying UV-B radiation intensities compared to that when wheat leaves were not treated with UV-B radiation. Overall, the effects of direct exposure of wheat plants to UV-B radiation with different intensities in different periods on epidemiological components of wheat stripe rust were systematically explored, and the results suggest that the effects of UV-B radiation increased gradually with the increase of UV-B radiation intensity. This information provides a basis for monitoring and predicting this disease as well as for conducting further studies on pathogen virulence variation.