国际上小麦叶锈菌生理小种鉴别寄主及其小种命名方法

由于地理生态条件、叶锈菌群体结构、研究历史和研究方法等的差异,不同国家和地区采用相应的小麦叶锈菌鉴别寄主和小种命名方法。美洲、中国、欧洲、埃及、南非等国家和地区均采用以Thatcher为遗传背景的分别含有Lr1、Lr2 a、Lr2 c、Lr3、Lr9、Lr16、Lr24、Lr26、Lr3 ka、Lr11、Lr17和Lr30基因的近等基因系或单基因系作为小麦叶锈菌鉴别寄主,但附加的辅助鉴别品种不尽相同;澳大拉西亚和印度的鉴别寄主自成体系。这种格局不利于国家间研究结果的对比和交流。因此,建立一套国际通用鉴别体系实属当务之急。     

中国小麦条锈菌鉴别寄主维尔对条锈病的温敏抗性研究

维尔是中国小麦条锈菌重要鉴别寄主,通过潜育期变温处理对其进行温敏抗性检测和基因推导分析。本研究发现,维尔对小麦条锈菌流行小种条中32号的抗性表现为侵染型由常温下的高侵染型向高温下的低侵染型转变,侵染指数降低,呈现高温诱导抗条锈性;通过与已知的微效基因品系比较及方差分析,推测维尔对条中32号的温敏抗性可能是由2对高温诱导表达的温敏微效基因控制。建议将维尔作为温敏微效基因资源在抗病育种中利用,作为鉴别寄主用于小种鉴定时,要严格控制鉴定温度,不要超过18 ℃。     

小麦条锈菌中国鉴别寄主阿勃抗条锈病基因遗传分析

阿勃是中国小麦条锈菌重要鉴别寄主之一,该研究采用经典遗传分析、单体分析等方法,通过不同条锈菌系鉴定,系统分析了阿勃抗条锈性的遗传基础及抗性特点.结果表明,在常温条件下,阿勃在苗期对中国小麦条锈菌系水源致病类型1(Su-1)和印度菌系79009的抗性均由2对抗条锈病基因控制,属细胞核遗传,抗79009茵系的2对基因分别定位在3B和7B染色体上,是与已知抗条锈病基因不同的未知新基因,分别暂定名为YrAbb1和YrAbb2.     

1997~2001年我国小麦条锈菌生理小种变化动态

 1997~2001年采自我国13个省和自治区的主要生产品种和重要抗源上的小麦条锈菌标样4008份,其鉴定结果基本反映了这期间我国小麦条锈菌生理小种的变化状况。1997年发现了水源11致病类型13和14,2001年把原Hybrid 46致病类型3(HY-3)命名为条中32号。这期间共监测到41个小种和致病类型,每年都能监测到的小种和致病类型有32个,为条中17、19、21、22、23、25、26、28号,洛10-2,条中29,洛13-2、3,条中30、31、32,Hybrid 46-4、5、6、7、8,水源11-2、3、4、5、6、7、8、10、11、12、13和14。条中29、30、31、32号为主要流行小种。1997~2000年间条中31号位居第一,出现频率为10.8%~13.5%,其次为条中32号(4.8%~11.7%),由于条中32号逐年持续上升,2001年超过条中31号而位居第一。其次是条中29和30号,两者出现频率基本相当,1997~1999年条中29号略高位第3(3.3%~4.0%),条中30号位第4(2.6%~3.3%),2000~2001年,条中30号略高位第3(3.6%~7.3%),条中29号位第4(2.7%~3.1%)。出现频率较高的还有条中23号、25号、26号、28号和Hybrid 46-4、5、7、8,水源11-3、4、5、7、11、13和14,其它为稀有小种和致病类型。相对于东部黄淮海流域,甘肃、四川、陕南和云南及其毗邻地区条锈菌群体结构复杂,内部多样性高,几乎所有小种和致病类型都可监测到。从整个条锈菌群体衡量Hybrid 46和水源11致病类群占有很大份额,1997~2001年两者总的年出现频率分别为78.5%、80.2%、76.6%、78.7%和87.4%,致使我国小麦条锈菌群体均匀度低,这与我国小麦品种抗病基因较简单、且主要集中在Yr9为重要抗源的"洛类"衍生系(80%以上)和以繁6为抗源的繁6衍生系有一定关系。     

中国小麦条锈菌鉴别寄主维尔抗条锈性遗传分析

小麦品种维尔是中国小麦条锈菌重要鉴别寄主.将该品种分别与完全感病品种铭贤169和其它抗病品种杂交,获得各组合的F1、BC1和F2代群体及部分F3家系.在温室对各组合亲本及F1、BC1和F2代群体进行了苗期抗性鉴定.结果表明,维尔对CY17和CY23菌系的抗性由1对主效隐性基因控制.等位性分析表明,维尔中抗CY17和CY23菌系的基因与Triticum spelta album中的1对抗性基因等位或紧密连锁,将其命名为YrVirl.而维尔对Su-1的抗性则由1对显性基因控制,将其命名YrVir2.同时采用CY17和CY23菌系对铭贤169/维尔组合的48个F3家系进行了苗期抗性鉴定,根据平均抗性指数和样本方差将这些家系分为3组,卡方测验证明了维尔对上述两菌系的抗性由1对主效基因控制,同时还可能受微效基因的影响.     

中国小麦条锈菌生理小种同工酶分析

 本文首次报道利用聚丙烯酰胺凝胶电泳技术分析我国小麦条锈菌11个主要流行生理小种及美国小麦条锈菌白化菌系夏孢子提取物中酯酶、过氧化物酶、过氧化氢酶、碱性磷酸酶、酸性磷酸酶、谷草酰胺转氨酶、多酚氧化酶的同工酶图谱。结果表明:我国小麦条锈菌不同生理小种具有同工酶表型异质性,而同一生理小种的不同单孢系间或不同地理来源的菌系间同工酶表型相同,即生理小种群体内部可能具有同工酶表型同质性。依据同工酶资料的统计分析结果,将所测的中国小麦条锈菌的11个生理小种划为5个类群:A群(条中8号)、B群(条中10号)、C群(条中17号)、D群(条中19、22、23、25、27、26号)、E群(条中28、29号),它反映出小种间遗传基础的相似性及演化特点。结合对美国条锈菌菌系同工酶测定,并与国外同类研究结果比较,揭示出中国小麦条锈菌具有独特的同工酶表型。同工酶表型分析是研究锈菌毒性变异的一个有力工具,建议建立起中国小麦条锈菌毒性菌系多种同工酶标准图谱,用作研究小麦条锈菌毒性变异和群体遗传学的参照系。     

1991年小麦条锈菌生理小种监测结果简报

 1991年中国农科院植保所与陕西、甘肃、四川、山西、云南省农科院植保所协作,监测了采自晋、冀鲁、豫、陕、甘、川、青、宁、滇、黔、皖、鄂、浙等14省(区)的共1049个小麦条锈菌标样。     

中国条锈菌新小种条中30、31号的研究

本文报道了1991年以来对新小种条中30、31号鉴定与致病性的研究。继1991年发现对绵阳系成株小麦有致病力的、对Hybrid46有毒的新致病类型91—1,1993年又发现了新的致病类型93—1。根据它们对我国鉴别寄主的反应,命名为条中30、31号。与条中28、29号相比,新小种具有更宽毒性基因组成和更高的相对寄生适合度值,它们对我国生产品种、高代品系和重要抗源有更广的致病范围。证实两个新小种的出现和发展是绵阳系小麦抗条锈变异的主要因素,建议加强对新小种抗病育种和流行预测的研究。     

小麦条锈菌鉴别寄主抗引655抗条锈性遗传分析

小麦品种抗引655是中国小麦条锈菌重要鉴别寄主。将该品种分别与完全感病品种铭贤169及其它抗病品种杂交，获各组合的F1、BC1和F2代群体。在温室对各组舍亲本及F1、BC1和F2代群体进行了苗期抗性鉴定。结果表明，抗引655对CY29菌系的抗性由2对显性抗条锈基因互补控制，对CY31和Su-1的抗性由1对显性基因控制。等位性分析表明，抗引655中抗CY29、CY31和Su-1的基因与Triticum speha album中的1对基因等位或紧密连锁。故而判定抗引655中除含有YrI外，还含有2对抗条锈基因，暂定名为YrKy1和YrKy2；YrKy1只抗CY29，YrKy2同时抗CY31和Su-1。系谱分析表明，抗引655中的抗条锈基因极有可能来自前苏联地方品种选育而成的旱熟36。     

小麦条锈菌鉴别寄主尤皮Ⅱ号抗条锈性遗传分析

小麦品种尤皮Ⅱ号是重要的中国小麦条锈菌鉴别寄主.为研究尤皮Ⅱ号的抗条锈性遗传基础,将该品种分别与感病品种铭贤169及其它抗病品种杂交,获得各组合的F1、BC1和F2代群体.在温室对各组合亲本及F1、BC1和F2代群体进行了苗期抗性鉴定.结果表明,尤皮Ⅱ号对CY29菌系的抗性由两对隐性基因独立或重叠遗传控制;对CY23的抗性由两对显性基因互补遗传控制;对CY31的抗性亦由两对显性基因互补遗传控制,而对Su-1的抗性则由一对显性基因控制.抗CY29的两对基因不抗CY23、CY31和Su-1,因此将这两对基因暂定名为YrJu1和YrJu2.抗CY23的两对基因中,其中一对同时抗CY31和Su-1,该基因与Spaldings prolific中的一对基因等位或紧密连锁,将该基因暂定名为YrJu3;另一对则与抗引655中的一对抗性基因等位或紧密连锁,暂定名为YrJu4.YrJu1、YrJu2、YrJu3和YrJu4均与其它供试品种中的已知抗性基因不同.     

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.     

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.     

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.     

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.     

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.     

Genetic analyses of resistance of the wheat differential cultivars Carstens V and Spaldings Prolific to two races of Puccinia striiformis

Genetic analysis of resistance of wheat seedlings to two races of Puccinia striiformis was conducted on F₁, F₂ and F₃ generations from crosses Carstens V (CV) × Lee, Spaldings Prolific (SPA) × Lee and CV × SPA. F₂ generations from crosses of CV and SPA with Strubes Dickkopf (SD) were also studied. The plants were classified into six resistance classes and analysed by factorial correspondence analysis and nonhierarchical classification. The two P. striiformis isolates tested were a French isolate of race 43E138 and a Lebanese isolate of race 2E16, selected for the differences in their virulence spectra for the common differential cultivars Strubes Dickkopf and Nord Desprez. Resistance of CV and SPA was recessive and dominant to races 43E138 and 2E16, respectively. CV possessed three or four resistance genes, one of them being expressed with both races. Two genes of CV had a cumulative effect for resistance to 43E138 and two or three gave dominant resistance to 2E16. SPA had three resistance genes, all of which gave resistance to 2E16 and two of which also gave resistance to 43E138. SPA had one gene in common with CV for resistance to both races. Furthermore, the gene for resistance to race 2E16 in CV and SPA was allelic with a gene in SD, and was probably Yr25 .     

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.     

2000～2001年我国小麦条锈病发生和生理小种监测结果

2000年全国小麦条锈病发生为非流行年 ,四川省属于中等偏重发生 ,其他麦区轻或零星发生。2001年全国小麦条锈病为中度流行年份 ,四川、云南和贵州为中度偏重流行 ,陕西、湖北和河南中部中度流行 ,甘肃中度偏轻发生 ,河南、山西、山东和河北等东部麦区轻和局部零星发生。对 14 82个小麦条锈菌标样鉴定结果表明 ,条中 32号(Hybrid 4 6类型3,HY-3)近年出现频率持续上升,2001年其出现频率首次居各小种及致病类型首位,为28.79%,预测今后一段时间内将成为我国的主要流行小种之一。条中31号2001年出现频率为9.51%,比2000年13.5%有所下降 ,居第2位。条中30号2001年出现频率为7.33%,比2000年3.55%有所回升 ,在各小种及致病类型中居第3位。抗条锈育种应以条中32、31、30号和水源11类型14为主要对象 ,兼顾条中29号、Hybrid46和水源11的其他致病类型     

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.