Evidence of host‐range expansion from new powdery mildew (Blumeria graminis) infections of triticale (×Triticosecale) in France

This study aimed to determine whether powdery mildew caused by Blumeria graminis is an endemic pathogen of triticale (×Triticosecale: Triticum × Secale), emerging as a result of recent changes in its pathogenicity, or whether it is a new pathogen, possibly resulting from hybridization between ff. spp. tritici and secalis. A secondary aim was to consider breeding practices that may have favoured this emergence. Phylogenetic analyses based upon six genes revealed the close relatedness of the novel entity and the ff. spp. tritici and secalis, but the IGS marker finally grouped together the isolates collected on triticale and on wheat, supporting the scenario of a recent host‐range expansion from wheat to triticale. Pathotype analyses concluded that virulence spectra of B. graminis infecting triticale were new in comparison to those observed for other reference formae speciales, and lack of fungicide resistance in triticale isolates strengthens the hypothesis of no or little genetic exchange between wheat and triticale populations of powdery mildew. This adaptation may follow the breakdown of plant resistance genes, which are probably not very diverse in current triticale cultivars since this criterion was not considered as a major one until recent years. Moreover, the complex selection and genetics of this hybrid cereal makes it difficult to predict the transmission of powdery mildew resistance genes.     

普通小麦“兰考90(6)”品系对白粉病抗性的遗传研究

 普通小麦(Triticum aestivum L.)"兰考90(6)"系列品系是以六倍体小黑麦(X Triticosecale Wittmack;AABBRR)为白粉病抗源培育的新的小麦-黑麦1BL/1RS异易位系。这些品系高抗白粉病。小麦白粉病抗性基因推导试验证明,"豫麦66"携带的抗病基因与大多数已经报道的小麦抗白粉病基因不同。用白粉菌[Blumeria graminis (DC.) E. O. Speer f. sp. tritici]单孢堆分离物进行的遗传分析表明,"兰考90(6)"品系携带一个小种专化的隐性抗白粉病基因。对"中国春"和"兰考90(6)21-12"杂交F₂分离群体进行1RS染色体检测,结果证明该抗白粉病基因不在1RS染色体臂上。本研究为有效利用"兰考90(6)"系列品系中的抗白粉病基因提供了科学依据。     

What types of powdery mildew can infect wheat-barley introgression lines?

This work is a detailed study of the infection of fungal biotrophic pathogens causing powdery mildew diseases on introgression lines originating from the intergeneric hybridisation between wheat and barley (Triticum aestivum L. × Hordeum vulgare L.). Powdery mildew fungi are among the most widespread biotrophic pathogens of plants also and infect dicot and monocot species. Most powdery mildew species are strictly host specific. They colonize only a narrow range of species or one particular host species. The intergeneric hybridisation between wheat and barley could result in expansions of host ranges of the barley powdery mildew. Our experiments covered natural infections in the field and artificial infections under greenhouse conditions. Formae speciales of powdery mildew were identified on the basis of the sequencing results of ribosomal internal transcribed spacer sequences (rDNA-ITS). We identified Blumeria graminis f.sp. tritici isolate 14 (HM484334) on the wheat parent and all wheat-barley introgression lines and B. g. f. sp. hordei isolate MUMH1723 (AB 273556) on the barley parent, respectively. The wheat-barley introgression lines were inoculated with barley powdery mildew under greenhouse conditions. According to our results the added barley chromosomes (or segments) do not cause host range expansion of barley powdery mildew.     

Virulence phenotypes in powdery mildew (Blumeria graminis) populations and resistance genes in triticale (x Triticosecale)

Triticale (xTriticosecale) is a new host for powdery mildew (Blumeria graminis) being not infected by this pathogen in Germany before 2001. To evaluate population structure of the pathogen and race-specific resistances in the host, 694 isolates were collected in 12 states of Germany in the years 2007 to 2009 on triticale. They were tested by a newly developed initial differential set of 20 triticale cultivars. Corresponding virulences were found for all differentials except for cultivar Grenado. In total, 272 different virulence phenotypes (=pathotypes) were detected. The virulence complexity of the isolates ranged from 6 to 19 of 20 possible virulences with a mean of 15. In all years, a high level of diversity of the powdery mildew populations was observed with Simpson indices in the range 0.95 to 0.97. The distribution of the pathotypes was even across Germany with an Evenness index in the range 0.82 to 0.88. A set of 19 isolates with different virulence pathotypes and 10 cultivars were selected to be used to identify race-specific resistances of triticale cultivars and breeding lines. Some cultivars susceptible to most of the isolates in seedling stage were moderately resistant in adult-plant stage. The high diversity and complexity of the pathotypes found in German powdery mildew populations as well as an increasing acreage of only a few dominating triticale cultivars accelerate the adaptation of the pathogen to race-specific host resistances suggesting restricted durability only. More durable resistance might be achieved by combining new effective race-specific (qualitative) resistance genes with race-nonspecific (quantitative) resistances effective in the adult-plant stage that are already available.     

小麦农家种红秃头和霸王鞭的抗白粉病基因分析及染色体定位

本研究对红秃头和霸王鞭两个小麦农家种抗白粉病基因推导显示,红秃头和霸王鞭均具有较宽的抗性谱,是良好的抗源品种,并可能携带新的抗病基因。抗白粉病遗传分析表明,红秃头对E09的抗性由1对显性基因控制,对E26和E30-2的抗性分别由1对隐性基因控制,其至少携带一显一隐2对抗白粉病基因;霸王鞭对E09的抗性由2对显性基因重叠或者独立控制,对E26的抗性由2对显性基因互补作用控制,对E30-2的抗性由1对显性基因控制,其至少携带2对显性基因。利用基因芯片结合集群分离分析法(Bulk Segregant Analysis,BSA)进行染色体定位推测出,红秃头的抗白粉病基因可能位于染色体7B和6B上,霸王鞭的抗白粉病基因可能位于染色体4A和7B上。     

三个小麦农家品种的苗期抗白粉病遗传分析

[目的]对3份小麦农家品种‘矮秆芒麦’、‘红头麦’和‘大红头’进行苗期抗性的遗传分析,研究它们的抗白粉病遗传特点,为其在抗病育种中的有效利用提供依据.[方法]将这3份小麦农家品种分别与感病品种‘铭贤169’正、反杂交,获得了F1和F2代.利用白粉菌E09菌株,分别对这3份农家品种、感病亲本‘铭贤169’以及各自的F1和F2代植株进行抗性鉴定.调查统计的数据经卡方测验分析其符合度.[结果]这3份农家品种对白粉菌E09菌株的抗性均由1对隐性核基因控制.[结论]3份农家品种对石家庄本地区的混合白粉病菌表现出良好的抗性,并且对E09的抗性均由1对隐性基因控制.可以进一步对它们进行分子标记及定位研究,为其作为抗源在小麦抗白粉病育种中的应用奠定基础.     

Comparative analysis of early H2O2 accumulation in compatible and incompatible wheat–powdery mildew interactions

Comparative analysis was carried out for hydrogen peroxide (H₂O₂) accumulation in response to powdery mildew (Blumeria graminis[Erysiphe graminis] f.sp. tritici) primary germ tube (PGT) and appressorial germ tube (AGT) contact, and in papilla (cell wall apposition) and hypersensitive responses (HR) in wheat (Triticum aestivum). Using primary leaves of three susceptible wheat lines (Bainong 3217; Beijing 837; Jingshuang 16) and five resistant lines [Mardler (Pm2 + Pm6); Ulka/8 × Cc (Pm2); Mardler/7 × Bainong 3217 (Pm2); Pm16; Pm16/7 × Beijing 837 (Pm16)], early H₂O₂ accumulation between 10 and 48 h after inoculation was studied. Strong H₂O₂ accumulation was found in effective papillae and associated cytosolic vesicles in both susceptible and resistant wheat lines, suggesting the important role of H₂O₂ in effective papillae formed as a general plant defence against powdery mildew. High frequency of effective papilla formation was observed in all five resistant lines. Among resistant lines, hypersensitive cell death was detected earlier in Mardler and Ulka/8 × Cc than in Pm16 and Pm16/7 × Beijing 837. In all cases this was associated with H₂O₂ accumulation in attacked epidermal cells. Interestingly, penetration resistance but not HR appeared to be mediated by the Pm2 gene in the Mardler/7 × Bainong 3217 line, suggesting that Pm2 may govern an HR-independent defence pathway in this genetic background. As effective papillae and HR did not occur in the same cells, papilla deposition may be independent of the HR response, despite the fact that both defence mechanisms were associated with high H₂O₂ accumulation. In wheat powdery mildew-incompatible interactions, HR acts as a second line of defence to contain infection when the papilla defence fails.     

Gene expression profile and physiological and biochemical characterization of hexaploid wheat inoculated with Blumeria graminis f. sp. tritici

Wheat (Triticum aestivum L.) powdery mildew has direct effects on photosynthesis- and energy-related pathways. Because it is prevalent in Sichuan Province, China, knowledge concerning Pm40-mediated wheat powdery mildew resistance is important for understanding host–pathogen interactions. The aim of this study was to understand physiological changes during the host–pathogen interaction, including gene expression, photosynthetic and chlorophyll fluorescence parameters, chlorophyll content, and antioxidant activity, specific to the resistance response to powdery mildew. The Pm40-expressing L693 and Pm40-deficient L1095 wheat lines were employed for comparison analyses. Fifty-eight and 26 expressed sequence tags (ESTs) were obtained from forward and reverse suppression subtractive hybridization cDNA libraries constructed from Bgt-infected L693 and L1095 leaves, respectively, and 69% of the ESTs in the forward library were related to photosynthesis- and energy metabolism-related pathways. Our data indicate that photosynthesis and carbon metabolism as well as nitrogen metabolism related to photosynthesis are the main pathways involved in Pm40-mediated wheat powdery mildew resistance and that an acetyl CoA-like gene might be a key regulatory factor in these pathways. This study provides new insight into the physiological and genetic mechanisms of resistance to wheat powdery mildew.     

细胞乳突的形成和小麦白粉菌成功侵染的关系

 用一套己知抗白粉病单基因的小麦材料,研究了小麦白粉菌诱发寄主表皮细胞产生的乳突与成功侵染的关系。抗病小麦叶片上,诱发产生乳突的分生孢子占测定分生孢子数的58.8%,感病品种为63.8%。这表明抗性不同的小麦材料接种白粉菌之后,叶表皮细胞内形成乳突的百分率相近,然从在抗病材料和感病材料中形成的乳突阻止病原物发育和成功侵染的作用不相同,在抗病叶片上,诱发产生乳突的分生孢于中有88%停留在压力孢阶段,不继续发育,仅有12%能突破乳突形成吸器,使得侵染成功。与此对照,在感病叶片上只有32%诱发乳突的分生孢子不能穿透乳突,而68%则能突破乳突。成功地与寄主建立寄生关系。我们的研究表明,乳突能否成功地阻止白粉菌侵入可能与乳突形成的迟早有关。     

A major gene for powdery mildew resistance transferred to common wheat from wild einkorn wheat

ABSTRACT A major gene for resistance to wheat powdery mildew (Blumeria graminis f. sp. tritici = Erysiphe graminis f. sp. tritici) has been successfully transferred into hexaploid common wheat (Triticum aestivum, 2n = 6x = 42, AABBDD) from wild einkorn wheat (Triticum monococcum subsp. aegilopoides, 2n = 2x = 14, AA). NC96BGTA5 is a germ plasm line with the pedigree Saluda x 3/PI427662. The response patterns for powdery mildew resistance in NC96BGTA5 were tested with 30 differential isolates of B. graminis f. sp. tritici, and the line was resistant to all tested isolates. The analyses of P(1), P(2), F(1), F(2), and BC(1)F(1) populations derived from NC96BGTA5 revealed two genes for wheat powdery mildew resistance in the NC96BGTA5 line. One gene, Pm3a, was from its recurrent parent Saluda, and the second was a new gene introgressed from wild einkorn wheat. The gene was determined to be different from Pm1 to Pm21 by gene-for-gene and pedigree analyses. The new gene was identified as linked to the Pm3a gene based on the F(2) and BC(1)F(1) populations derived from a cross between NC96BGTA5 and a susceptible cultivar NK-Coker 68-15, and the data indicated that the gene was located on chromosome 1A. It is proposed that this new gene be designated Pm25 for wheat powdery mildew resistance in NC96BGTA5. Three random amplified polymorphic DNA markers, OPX06(1050), OPAG04(950), and OPAI14(600), were found to be linked to this new gene.     

Selection for increased spore efficacy by host genetic background in a wheat powdery mildew population

ABSTRACT A field experiment was designed to test the hypothesis of for increased reproductive ability on different host genetic backgrounds within a wheat powdery mildew population. Studies have suggested that, in host mixtures, such selection could increase the reproduction rate of simple patho-types that always develop on the same host genetic background, whereas complex pathotypes should not be affected because they infect different host genotypes. In our experiment, the Erysiphe graminis population reproduced for successive generations on cvs. Orkis and Etecho, either grown as pure stands or in a mixture. In an additional treatment, the host cultivar changed after each generation. Isolates were sampled in April and, after seven pathogen generations, in July. At the second sampling date and for pure stands only, mean spore efficacy was greater on the host from which isolates were sampled than on the other one. This was attributed to selection within the pathogen population for better spore efficacy on the host genetic background. This selection was independent of the virulence genes carried by the isolates. The possibility of a phenotypic plasticity effect was tested and rejected.     

Differential effects of phenylalanine ammonia lyase, cinnamyl alcohol dehydrogenase, and energetic metabolism inhibition on resistance of appropriate host and nonhost cereal-rust interactions

ABSTRACT Effects of phenylpropanoid and energetic metabolism inhibition on resistance were studied during appropriate host and nonhost cereal-rust interactions. In the appropriate barley-Puccinia hordei interaction, phenylalanine ammonia lyase (PAL) and cinnamyl alcohol dehydrogenase (CAD) inhibition reduced penetration resistance in two genotypes, suggesting a role for phenolics and lignins in resistance. Interestingly, penetration resistance of the barley genotype 17.5.16 was not affected by phenylpropanoid biosynthesis but penetration resistance was almost completely inhibited by D-mannose, which reduces the energy available in plant host cells. This suggests a parallel in the cellular basis of penetration resistance between 17.5.16 rust and mlo barleys powdery mildew interaction. Results revealed differing patterns of programmed cell death (PCD) in appropriate versus nonhost rust interactions. PAL and CAD inhibitors reduced PCD (hypersensitivity) in appropriate interactions. Conversely, they had no effect in PCD of wheat to P. hordei; whereas D-mannose dramatically reduced nonhost resistance and allowed colony establishment. The differential effects of inhibitors in the expression of the different resistances and the commonalities with the cereal-powdery mildew interaction is analyzed and discussed.     

Resistance to powdery mildew (Blumeria graminis f.sp. avenae) in oat seedlings and adult plants

In this work, 165 Avena sativa and Avena byzantina accessions were screened for resistance to powdery mildew caused by Blumeria graminis f.sp. avenae and the defence mechanisms of resistant plants were further characterized. Ten resistant and moderately resistant accessions were selected according to macroscopic assessment. A detailed histological study of selected genotypes showed a range of defence mechanisms, acting alone or in combination, that impeded fungal development at different stages. Since the resistance observed in the collection was scarce, a study of adult plant resistance was carried out in 45 genotypes selected from field trials. Nine oat landraces and two commercial varieties showed very high levels of adult plant resistance. A detailed study of the components of the adult plant resistance revealed a high increase of penetration and post‐haustorial resistance in the fifth compared to the first leaves. Identification of the resistance sources and characterization of underlying defence mechanisms will be useful for future breeding programmes and for further cellular and molecular studies to unravel the genetic basis of resistance, in this species in particular and in cereal–powdery mildew interactions in general.     

小麦-粗山羊草双二倍体抗白粉病基因定位及其遗传转移

中国春单体分析表明,四倍体小麦－粗山羊草双二倍体Ａｍ６的抗白粉病基因位于１Ａ染色体上,与２４个已知抗白粉病基因比较说明,Ａｍ６可能为一新的抗白粉病基因。限制性片段长度多态性分析表明,在分布小麦７个部分同源群的３６个探针中未找到与此抗病基因连锁的分子标记。     

小麦抗白粉病品种的基因分析与归类研究

 研究出用抗谱及抗谱的近似,分析小麦抗白粉病品种的基因与归类的方法,测试了152个小麦抗白粉病品种,归类出15个类群,共120个品种,分析了类群主效抗病基因,并根据测试结果分析提出,为聚合主效抗病基因,进行复合杂交,在选取抗病亲本对上,要掌握毒性频率低,抗谱上互补,抗病基因非等位三项基本原则,才能取得预期成效。抗谱、近似分析法快速,准确,能同时测量大量品种、获得较多可靠信息,适用于基因对基因专化性病害抗病品种的基因分析与归类。     

Interaction between powdery mildew and barley with mlo5 mildew resistance

Powdery mildew infection of barley with the mlo5 barley powdery mildew resistance gene was examined, using near-isogenic barley lines, with and without mlo5 resistance, and two near-isogenic powdery mildew isolates, HL3/5 and GE3 with high (virulent) or low (avirulent) penetration efficiency on the resistant barley line. In all isolate–host combinations (except GE3 on the resistant barley line), frequency of haustorium formation increased significantly from zero at 11 h after inoculation to a maximum by 13 h, and there was no subsequent increase up to 24 h. In the susceptible barley line, 27% of appressoria from both isolates formed haustoria. Although this was significantly higher than the frequency of haustorium formation (18%) of HL3/5 on the resistant barley line, HL3/5 was much more successful than GE3 (frequency of haustorium formation less than 1%). The fact that HL3/5 did not possess a generally higher ability to penetrate successfully to form haustoria on the susceptible barley line, indicates that HL3/5 did not overcome the mlo5 resistance by being generally more vigorous. In the resistant barley line, papillae were larger than in the susceptible line; however, both isolates were associated with papillae of the same diameter at the time of penetration. We suggest that the mlo5 resistant barley line confers two different forms of resistance: isolate-specific and isolate-nonspecific.     

36个小麦审定和区试品种(系)的抗白粉病基因推导

明确我国当前小麦审定和区试品种含有的抗白粉病基因, 可为这些品种在小麦抗病育种中的应用及品种合理布局和轮换提供依据。本研究采用24个不同毒性的小麦白粉菌菌株对36个小麦审定和区试品种(系)进行抗白粉病基因推导, 参试品种(系)与46个已知抗病基因小麦品种(系)抗谱比较的结果表明, 11个小麦审定品种中有5个品种对所有供试白粉菌菌株表现抗性, 结合亲本溯源, 推测其中有3个品种可能携有Pm21基因; 另外6个审定品种中有5个品种可能含有抗病基因Pm2, 且其对应的亲本或亲本组合中含有抗病基因Pm2。25个区试品种(系)中有3个可能含有Pm21, 10个含有Pm2, 1个含有Pm2+6,2个含有Pm4b,1个含有Pm8。另外, 参试的36个品种(系)中还有9个品种(系)和已知基因品种抗谱存在一定差异。总体上, 推导出已知基因的品种以含有Pm2基因的品种最多, Pm21基因的品种次之, 建议在生产上加强对Pm21基因品种(系)特别是已审定的携有Pm21基因品种的推广和应用, 应该注意一些省份在育种和生产上应慎用或少用含Pm2基因的品种(系)。     

2002年部分麦区小麦白粉病菌对三唑酮的抗药性监测及苯氧菌酯敏感基线的建立

 采用拌种离体叶段法测定了2002年采自北京、江苏、山东、山西、河北、新疆和四川7省市部分麦区的109个小麦白粉病菌菌株对三唑酮和苯氧菌酯的敏感性,结果表明小麦白粉病菌群体对三唑酮敏感性EC₅₀平均值是63.70 μg/mL,平均抗性水平为30.48倍,最高抗性水平达142.97倍。其中江苏、山东、四川三地的菌株抗药性水平明显高于河北、北京、山西和新疆。同时测得病菌群体对苯氧菌酯的敏感基线EC₅₀值为85.82μg/mL,且白粉病菌对甲氧基丙烯酸酯类杀菌剂苯氧菌酯和三唑类杀菌剂三唑酮之间无交互抗性。该结果可为这两类杀菌剂在生产上的合理应用和抗药性治理提供依据。     

小麦白粉菌有性时期闭囊壳在侵染循环中作用的初步研究

通过对陕西省和河南省小麦白粉病越夏期间场院麦秸垛中闭囊壳发育、存活情况及场院附近自生麦苗发病情况的调查,结果发现闭囊壳保湿培养7~8 d后,子囊孢子形成达到高峰;自生麦苗发病率与闭囊壳存活率之间存在极显著的正相关性,且闭囊壳存活率与海拔高度之间存在显著的负相关性,采自海拔500~900 m的闭囊壳有较高的存活率,海拔超过1 000 m的存活率较低或为零。此结果说明小麦白粉菌有性时期闭囊壳可能是通过侵染自生麦苗在侵染循环中起作用。