Local dispersal of Puccinia striiformis f. sp. tritici from isolated source lesions

Understanding how disease foci arise from single source lesions has not been well studied. Here, single wheat leaves were inoculated with Puccinia striiformis f. sp. tritici urediniospores, and all wheat leaves within two intersecting 0.3 × 3.0 m transects were sampled in eight replicates over 3 years. The lesions observed on each of the top three leaves on plants within 1.5 m from the source lesion were three‐dimensionally mapped. The total number of lesions within a 1.5 m radius was estimated by dividing the number of lesions observed within each 0.025 m‐wide annulus by the fraction of the annulus sampled. The estimated total number of lesions produced within 1.5 m of a single source lesion ranged from 27 to 776, with a mean of 288 lesions. Eighty percent of the lesions were recorded within 0.69 m of the source infection. The proportion of total lesions observed at a given distance from the source was fitted well by the Lomax and Weibull distributions, reflecting the large proportion of lesions arising close to the source, and when fitted by an inverse power distribution had a slope (b) of 2.5. There were more lesions produced on leaves higher in the canopy than on lower leaves, with more lesions being detected above than below the point of inoculation. Simultaneous measurement of lesion gradients and spore dispersal in the final year of the study suggests that this pattern is due to greater susceptibility of upper leaves, rather than increased dispersal to upper leaves.     

Thermal generalist behaviour of invasive Puccinia striiformis f. sp. tritici strains under current and future climate conditions

Yellow rust is a devastating wheat disease. Since 2000, Puccinia striiformis f. sp. tritici strains PstS1 and PstS2 have become adapted to high temperatures and have spread worldwide. By 2011, Warrior strains had invaded both warm and cold areas of Europe. This study questioned whether thermal aptitude promoted the spread of Warrior strains, similar to PstS1/PstS2, by comparing infection efficiency (IE) at five temperatures and latent period (LP) under warm and cold regimes for Warrior isolates and pre‐2011 reference strains on two susceptible wheat varieties. The Warrior isolates showed a range of IE and LP responses to temperature that was intermediate between the northern reference isolates adapted to cold conditions and both the southern and invasive PstS2 isolates adapted to warm conditions. Warrior isolates had the highest IE under optimal temperatures of 10 and 15 °C, and displayed reduced infectivity under the warmest (20 °C) and coldest (5 °C) temperatures. Warrior strains acted as thermal generalists and the reference isolates acted as specialists. An IE thermal response was used to simulate the development of each isolate under future climate scenarios in a temperate and Mediterranean region. Isolates had the same ranking for yearly IE over the three 30‐year periods (1971–2000, 2021–2050, 2071–2100) and both locations, with a slight infection increase in the future. However, in the future IEs increased in earlier months. The thermal generalist profile of Warrior isolates for IE was confirmed, with an intermediate capacity to tolerate warming climate, whereas the southern isolates are better adapted to warm conditions, but do not have the virulences necessary to develop on current varieties.     

Clonality and long-distance migration of Puccinia striiformis f.sp. tritici in north-west Europe

The biotrophic fungus Puccinia striiformis f.sp. tritici , a basidiomycete that causes yellow rust on wheat, is spread by wind-dispersed spores. Analysis of amplified fragment length polymorphism (AFLP) variation showed that the fungus frequently migrates between the UK, Germany, France and Denmark. There is no biological evidence for sexual or parasexual reproduction under natural conditions, and this was supported by the lack of recombination, as revealed by AFLP, over the time and area represented by the samples in this study. A phylogeographic analysis revealed that there was effectively a single, clonal population in the four countries, up to 1700 km apart, consistent with a 'continent-island' model in which Denmark is the recipient of migrants from other countries. In five cases, specific pathogen clones were dispersed between the UK and Denmark, and on at least two recent occasions clones were also spread from the UK to Germany and France, causing outbreaks of yellow rust on wheat cultivars that were previously resistant to the disease in these countries. The agronomic consequences of migration were enhanced because of the limited genetic diversity for yellow rust resistance in wheat cultivars in the area. These results demonstrate that long-distance migration of pathogen clones, coupled with low diversity in the host species, may cause previously useful resistance genes to become ineffective for disease control on a continental scale.     

Support for a stepwise mutation model for pathogen evolution in Australasian Puccinia striiformis f.sp. tritici by use of molecular markers

Since its initial detection in Australia in 1979, wheat yellow (stripe) rust ( Puccinia striiformis f.sp. tritici ) has evolved in Australia and New Zealand into more than 20 pathotypes with assorted virulence characteristics. This evolution is believed to have occurred in a stepwise fashion from an original single pathotype, with no subsequent new introductions. A combination of random amplified polymorphic DNA (RAPDs) and amplified fragment length polymorphisms (AFLPs) was used to examine the level of molecular variation in Australian and New Zealand isolates, and to compare this with variation amongst other isolates of P. striiformis . Using 60 RAPD primers on seven Australian isolates representing seven different pathotypes collected between 1979 and 1991, more than 300 potentially polymorphic loci were analysed and no polymorphisms were detected. Using the same primers on two UK isolates, 3% of loci showed a polymorphism. A similar level of polymorphism was found between UK isolates using AFLP primers, and between 5 and 15% of fragments were polymorphic between an isolate from the UK, an isolate from Denmark, and one from Colombia. However, no AFLP polymorphisms were found amongst 14 Australian and New Zealand isolates tested, at over 100 potentially polymorphic loci. The lack of molecular variation in the Australian and New Zealand collection is consistent with the stepwise mutation theory of pathotype evolution from a single introduction.     

The recent history of Puccinia striiformis f.sp. tritici in Denmark as revealed by disease incidence and AFLP markers

Disease observations and amplified fragment length polymorphism (AFLP) markers were used to study recent developments in the Puccinia striiformis f.sp. tritici population in Denmark. The fungus appeared spontaneously at 10 locations in Denmark in 1997 after it was not observed under natural conditions in 1996. The pattern of disease development and prevailing winds suggested that the fungus reappeared by airborne spores from the south or west. In 1998, disease incidence was more evenly distributed throughout the country. Forty-eight single lesion isolates were collected from most crops where the disease was observed in these years; all except one from 1997 belonged to two pathotypes that were not previously detected in the country, and both possessed the newly discovered Yr17 virulence. The isolates were characterized with AFLP markers together with 28 isolates representing eight of 13 pathotypes observed prior to 1996. Initial screening of 240 Pst I/ Mse I AFLP primer combinations on four isolates showed that a primer combination, on average, revealed 0·4 polymorphisms between any isolate pair. A selection of 21 primer combinations resulted in 28 AFLP markers, which revealed 16 AFLP phenotypes among all 76 isolates. The two Yr17- virulent pathotypes consisted of three AFLP phenotypes, which were observed in both 1997 and 1998; the two most frequent AFLP phenotypes occurred at most sampling locations and often within the same crop. AFLP diversity was larger among samples collected prior to 1996, and also in this period most AFLP phenotypes were observed at different sampling locations. These results are consistent with the features of an entirely asexually reproducing pathogen dispersed by aerial spores across large areas.     

小麦条锈菌冬孢子生物学特性研究进展

由小麦条锈菌引起的小麦条锈病是我国广大麦区的严重病害,研究其冬孢子产生原因与作用对于解析该病菌的生活史及遗传变异机制至关重要.通过对国内外相关研究成果的整理,综述了小麦条锈菌冬孢子的形态特征、生物学特性、寄主范围及其作用与功能等方面的研究进展.     

Disease severity and pathotype dynamics of Puccinia striiformis f.sp. tritici in Denmark

Disease severity of wheat yellow rust, Puccinia striiformis f.sp. tritici , was analysed in Denmark from 1985 to 1999 in relation to the effects of weather on winter survival, distribution of host cultivars and pathotype dynamics. Below-average temperatures in January and February (midwinter) reduced yellow rust on the susceptible cv. Anja, and in three of four growth seasons following cold winters no yellow rust was observed on any cultivar under natural conditions. The agronomic consequences of dispersal of yellow rust urediniospores from external sources to Denmark, in a period during which large areas were planted with relatively few wheat cultivars, were demonstrated in several cases, most evidently when the Yr9 and Yr17 resistance genes became ineffective. Yr9 was overcome by the pathogen in a period with severe yellow rust epidemics on commercial cultivars, while virulence for Yr17 was first observed in a year with almost no yellow rust. In contrast, the resistance in cv. Kraka ( Yr1, CV ) was increasingly effective in controlling yellow rust, because pathotypes with the matching combination of virulence declined in the pathogen population. Pathotype frequency dynamics were thus influenced by selection forces within the country, and by selection forces in areas where spores were spread to Denmark from outside. The importance of a sufficient level of partial resistance in the wheat germplasm to prevent too much damage by yellow rust epidemics, in the event that the resistance genes are overcome by the pathogen population, is emphasized.     

Virulence analysis of Puccinia graminis f.sp. tritici populations in Ethiopia with special consideration of Ug99

Wheat stem rust samples were collected in 2006 and 2007 in the Arsi, Bale, Shewa and northwest regions of Ethiopia to determine virulence diversity and race distribution in Puccinia graminis f.sp. tritici populations. Stem rust incidence was high in Arsi, Bale and east Shewa. In northwest Ethiopia, and north and west Shewa, stem rust was prevalent at low levels. A total of 152 isolates was analysed and 22 races were identified. Races TTKSR (Ug99), TTHSR and RRTTR were predominant, with frequencies of 26·6, 17·7 and 11·1%, respectively. These races were also detected in all regions. The highly virulent race designated Ug99 was present throughout the country and dominated in all regions except northwest Ethiopia. A variant of Ug99 virulent against the stem rust resistance gene Sr24 was not detected in this study. Four stem rust resistance genes ( Sr13, Sr30, Sr36 and SrTm p) were found to confer resistance to most of the races prevalent in Ethiopia. With the exception of Sr30 , which is not effective against Ug99, these genes could be used in breeding for resistance to stem rust in Ethiopia.     

陕西省小麦条锈菌群体遗传结构分析

 利用TP-M13-SSR自动荧光检测技术,对陕西省小麦条锈菌群体遗传结构进行了分析。研究结果显示,在物种水平上,陕西省小麦条锈菌群体Nei’s基因多样性指数（H）为0.18、Shannon 信息指数（I）为0.29,表明该省条锈菌群体遗传多样性较为丰富。同时,条锈菌群体遗传多样性在地区之间也存在明显的差异,在7个条锈菌群体中,宁强种群遗传多样性相对较高（H为0.18,I为0.28）。AMOVA分析显示,陕西省小麦条锈菌在地区间、种群间和群体内的遗传分化分别为：12.26%、10.88%和76.86%,表明陕西省小麦条锈菌群体存在一定的遗传分化,但遗传变异主要发生在群体内部。     

Success and failure of invasive races of plant pathogens: The case of Puccinia striiformis f. sp. tritici in France

Invasions of new races can have contrasting consequences on populations of Puccinia striiformis f. sp. tritici, causing yellow rust of wheat. For example, the emergence of PstS7 (Warrior race) had major impacts in Europe and in France. By contrast, PstS2 had no impact in France, while it significantly affected other parts of the world. The objective of this study was to better understand factors that govern the success of an invasive race, taking the contrasting history of PstS7 and PstS2 in France as a case study. We compared these two races for three key factors driving invasive potential: (a) virulence against local cultivars, (b) aggressiveness in local environmental conditions, and (c) competitiveness against local races. During the period when PstS2 was detected, 70% of the grown wheat area was protected against this race by at least one known Yr resistance gene. By contrast, we found that only 15% of the wheat area had a low risk of infection by PstS7. In planta competition experiments suggested a higher competitiveness of PstS7 against local isolates compared to PstS2 in optimal thermal conditions. In silico experiments, based on thermal performance curves, suggested a high competitiveness of PstS7 considering infection efficiency. PstS2 was extremely competitive against local races in all considered environments (20 French sites × 15 years) due to its short latency period. Our findings highlight the importance of considering adaptation to environmental conditions, particularly temperature, in addition to virulence spectrum, in order to understand the evolutionary trajectories of emerging strains in pathogen populations.     

Genetic analysis and molecular mapping of resistance to Puccinia striiformis f. sp. pseudo‐hordei in common wheat

This is the first genetic study reporting on the interaction and molecular mapping of resistance to the barley grass stripe rust pathogen (Puccinia striiformis f. sp. pseudo‐hordei, Psph) in common wheat. Seedlings of 638 wheat accessions were tested and it was determined that wheat is a near‐nonhost to Psph based on rare susceptibility observed in <2% of commercial cultivars and <5% of wheat landraces. As previously observed for P. striiformis f. sp. tritici (Pst), the Australian cultivar Teal was highly susceptible to Psph. In contrast, a selection of cv. Avocet carrying complementary resistance genes Yr73 and Yr74 (Avocet R; AvR) was resistant. The Teal × AvR (T/A) doubled haploid (DH) population was used to map resistance in AvR to Psph. Infection types on the T/A DH lines inoculated with Psph and Pst indicated that all DH lines carrying both Yr73 and Yr74 were also resistant to Psph; however, fewer DH lines were susceptible to Psph than expected, suggesting the resistance was more complex. QTL analysis using 9053 DArT‐Seq markers determined that resistance to Psph was polygenically inherited and mapped to chromosomes 3A, 3D, 4A and 5B. The 3DL and 5BL markers co‐located with Yr73 and Yr74, suggesting an overlap between host and non‐host resistance mechanisms.     

小麦条锈菌鉴别寄主和小种命名现状

 中国是世界上小麦条锈病最大的流行区之一,也是限制中国小麦生产的最严重病害之一,特别是当条件适于其发生时会造成巨大的产量损失.小麦条锈菌小种生理专化性研究及小种监测是掌握小麦条锈菌及其毒性变化动态、抗病基因有效性及病害流行预测预报的重要依据,是治理由此病原菌引起的小麦条锈病的基本环节之一.由于地域、条锈菌群体结构、研究水平和历史等的差异,目前国际上不同国家和地区采用相应的小麦条锈菌鉴别寄主和小种命名方法.中国、美国和印度均使用了各自不同的小麦条锈菌鉴别寄主,并采用了相应的小种命名体系,例如从1957至2002年,中国利用10多个鉴别寄主已鉴定和命名了32个小种和30多个致病类型,美国利用10多个鉴别寄主已鉴定和命名了80个小种;另一类是采用1972年Johnson提出的二进制国际小种命名法,这种体系包括了国际鉴别寄主和欧洲鉴别寄主,象一套鉴别寄主被整体使用,欧洲、中东、亚洲、非洲和南美国家及澳大利亚和新西兰均采用此体系监测小麦条锈菌变化,是一套被最为广泛接受和使用的小麦条锈菌小种鉴别方法和命名系统,但随着研究深入和各国实际情况差异等原因,不同研究人员加入了不同的辅助鉴别寄主,以更能准确和实际地反映本国和地区的现实情况,为生产服务,本文对此现状进行了分析和比较.由于此病菌具有长距离气传和扩散的特点(地区间、国家间和洲际间),因而也讨论了未来小麦条锈菌鉴别寄主组成、小种命名方法和研究结果的交流和比较等实际问题.     

Relationships and genetics of wheat effects on infection frequency and colony extension of Puccinia striiformis f. sp. tritici

Wheat cvs Aquileja and Xian Nong 4, previously reported to possess quantitative resistance to Puccinia striiformis f. sp. tritici (Pst), were crossed to assess resistance associated with infection frequency and colony extension of Pst. The parents, F ₁, F ₂, F ₃, F ₄, backcrossed to Aquileja, and backcrossed to Xian Nong 4 were sown in the field. Experimental plants were inoculated with a single-spore derived isolate of Pst. Penetration frequency, colony size and disease severity were quantified. Comparison between the parents indicated that Aquileja was better at limiting infection frequency of Pst, whereas Xian Nong 4 was better at restricting colony extension. Penetration frequency and colony size were controlled by different genes. It was estimated that three or four genes, and three genes, controlling penetration frequency and colony size respectively, segregated in the cross Aquileja × Xian Nong 4, and were transmitted with moderately high heritability in additive fashion with no detectable dominant and epistatic effects. The two traits were correlated with disease severity. Colony size and penetration frequency explained, respectively, up to 33 and 8% of the genotypic variation of disease severity. These results will be helpful in understanding quantitative resistance and breeding for enhanced resistance to Pst.     

Russian populations of Puccinia triticina in distant regions are not differentiated for virulence and molecular genotype

The objective of this study was to determine whether genetically distinct groups of Puccinia triticina are present in four regions of the Russian Federation. Collections of P. triticina were obtained from the Central, North Caucasus, Volga and West Siberia regions from 2006 to 2010. Ninety‐nine single uredinial isolates were tested for virulence phenotype with 20 Thatcher near‐isogenic lines of wheat. Forty‐one virulence phenotypes were found in the four regions, with eight in common between the widely separated Central and West Siberia regions. A total of 72 isolates were tested for molecular genotype with 23 simple sequence repeat (SSR) primer pairs, and 66 isolates were used for further analysis after clone correction for virulence and molecular genotype. Analysis of variation showed no overall differentiation of SSR genotypes or virulence phenotypes based on region of origin. Linkage disequilibria for SSR genotypes were high across the entire population. The regional populations had higher than expected levels of allelic heterozygosity that indicated clonal reproduction. Based on cluster analysis of SSR genotypes there were two groups of P. triticina isolates that were widely distributed across Russia. The two SSR groups also differed significantly for virulence. Puccinia triticina may be dispersed from a common source of inoculum in the European or Caucasus regions of Russia. The Russian P. triticina populations were highly differentiated for SSR genotype from populations in Tajikistan, Kyrgyzstan, Uzbekistan, Armenia, Georgia and Azerbaijan and more similar to populations from southern Kazakhstan and northern Kazakhstan.     

Genetic differentiation of the wheat leaf rust fungus Puccinia triticina in Europe

The objective of this study was to determine whether genetically differentiated groups of Puccinia triticina are present in Europe. In total, 133 isolates of P. triticina collected from western Europe, central Europe and Turkey were tested for virulence on 20 lines of wheat with single leaf rust resistance genes, and for molecular genotypes with 23 simple sequence repeat (SSR) markers. After removal of isolates with identical virulence and SSR genotype within countries, 121 isolates were retained for further analysis. Isolates were grouped based on SSR genotypes using a Bayesian approach and a genetic distance method. Both methods optimally placed the isolates into eight European (EU) groups of P. triticina SSR genotypes. Seven of the groups had virulence characteristics of isolates collected from common hexaploid wheat, and one of the groups had virulence characteristics of isolates from tetraploid durum wheat. There was a significant correlation between the SSR genotypes and virulence phenotypes of the isolates. All EU groups had observed values of heterozygosity greater than expected and significant fixation values, which indicated the clonal reproduction of urediniospores in the overall population. Linkage disequilibria for SSR genotypes were high across the entire population and within countries. The overall values of R_ST and F_ST were lower when isolates were grouped by country, which indicated the migration of isolates within Europe. The European population of P. triticina had higher levels of genetic differentiation compared to other continental populations.     

Nitrogen per unit leaf area affects the upper asymptote of Puccinia striiformis f.sp. tritici epidemics in winter wheat

Field trials tested which components of epidemic development of Puccinia striiformis , the cause of yellow rust, were affected by nitrogen (N) fertilizer applied to winter wheat. Both timing and amount of N were varied to affect canopy size and leaf N content, and to provide a supply of mobile N to the pathogen, by causing fresh N uptake after leaf expansion was complete. No N was applied to control plots. A logistic disease-progress function was fitted to disease-severity data, which were assessed in absolute units. Leaf area and specific leaf N (g N per m² leaf tissue) were quantified. Large and highly significant effects of N on the upper asymptotes, or 'carrying capacities' ( c ) were found. Effects on rates and points of inflection of the epidemics were not significant. Early N resulted in larger shoot numbers and leaf area, but disease was also more severe, so that by grain filling, the remaining green leaf areas were larger without N than with N. Later N treatments did not increase canopy size, but did increase symptom area compared with the control. These effects differ from the concept that N affects disease as a result of its effect on canopy growth, and therefore canopy microclimate, and suggest instead a substrate effect. Linear regression revealed that 51% of the observed differences in c were explained by variation in specific leaf N, suggesting that growth of the rust fungus may depend directly on particular components of total leaf N.     

Histological investigation of stripe rust (Puccinia striiformis f.sp. tritici) development in resistant and susceptible wheat cultivars

The wheat cultivar Kariega expresses complete adult plant resistance against stripe rust, whereas cv. Avocet S is susceptible. Using confocal laser scanning microscopy, initial fungal penetration into flag leaves was identical in both cultivars, with directional germ-tube growth towards stomata that were penetrated without the formation of an appressorium, followed by differentiation of a substomatal vesicle, infection hyphae, haustorial mother cells and haustoria. During the following 4 days, further fungal development occurred more quickly in the resistant than in the susceptible cultivar. However, by 7 days postinoculation (dpi) the situation changed, with exponential growth of the pathogen occurring only in the susceptible line. Induced cellular lignification, a typical defence reaction of cereals, was observed at 4 dpi in the resistant cultivar, and 2 days later lignified tissue completely surrounded the fungal colonies. In the susceptible cultivar, isolated lignified host cells occurred at 6 dpi, and long, unbranched fungal hyphae outgrowing the resistance reaction were observed.     

中国小麦条锈菌主要流行菌系的寄生适合度研究

 为了预测我国未来小麦条锈菌的优势种群,本研究采用抗性组分法测定了我国条锈菌主要流行菌系的寄生适合度。结果表明:决定我国小麦条锈菌最重要的寄生适合度属性为条锈菌的产孢能力(繁殖力),其次为条锈菌的侵染能力,再次为扩展能力;建立了小麦条锈菌寄生适合度属性3个主成份的数学模型;在供试的菌系中,Su11-4、Su11-14和CY32相对寄生适合度较高,已经成为我国目前小麦条锈菌的优势小种(类型);CY31由于其寄生适合度较低,出现频率逐年下降,已成为次要小种,感染"中四"苗期的新菌系T4类型,由于其寄生适合度低,也不会成为今后一个时期的流行小种。     

中国小麦条锈菌4个主要流行小种的寄生适合度

 以12个全国大面积种植的小麦生产品种和1个高感品种为试材,利用抗性组分法研究了中国小麦条锈菌(Puccinia striiformis f.sp.tritici,Pst)4个主要流行小种条中32号、条中31号、水源11-14和水源11-4的寄生适合度。因子分析结果表明,病害量是度量条锈菌生理小种寄生适合度最重要的参数,其次是夏孢子生活力、产孢能力、产孢面积、产孢期,由此提出寄生适合度=(产孢量×侵染概率×夏孢子堆密度×夏孢子萌发率)/潜育期。研究还发现,条中32号的寄生适合度最高,致病性最强,与目前该小种为优势小种的事实相符;水源11-14的寄生适合度较高,有较大的发展潜力,在抗病育种中应予以高度重视。     

林芝地区小麦条锈菌转主寄主小檗的鉴定与分布

西藏自治区是我国小麦条锈病发生与流行相对独立的一个区系,小麦条锈病是严重影响西藏小麦安全生产的重要病害之一,林芝地区是西藏自治区小麦条锈病重要的常发区和流行区。已有研究证实,小檗是小麦条锈菌的转主寄主之一,并在我国小麦条锈菌致病性变异产生新菌系和小麦条锈病的发生过程中起作用。林芝地区小檗种类较多,但目前对林芝地区小檗是否能作为小麦条锈菌的转主寄主缺乏相关的研究和报道。本文针对小麦条锈病常发区的西藏林芝地区小檗资源进行了调查,对小檗能否作为小麦条锈菌转主寄主进行了鉴定。结果表明6种小檗在林芝地区常见且分布广泛,均是小麦条锈菌的转主寄主。这对进一步研究西藏小麦条锈病的发生、流行与病原菌致变性变异起着重要作用。