小麦与锈菌互作中TaAbcl和Sr33相关基因的功能研究
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摘要
植物的过敏性坏死反应（hypersensitive response, HR），被认为是植物在侵染位点形成的一种快速的死亡方式其和植物对病原菌的防御反应直接相关。在植物中，HR通常是在寄主和相关病菌的非亲和互作中被诱发的。小麦条锈菌（Puccinia striiformis f. sp.tritici，Pst）导致的小麦条锈病是一个威胁小麦生产的世界性问题。因而研究小麦（Triticum aestivum L.）和条锈菌防御反应过程中的相关基因，对揭示小麦和条锈病互作中的HR的分子机制以及抗锈性的信号转导途径奠定理论基础和提供理论依据。
为了寻找小麦对条锈菌防御反应中的基因，我们克隆并鉴定了一个新的类似Abc1蛋白家族的基因。我们对小麦Abc1基因进行了条锈菌侵染的生物胁迫和非生物胁迫以及小麦防御其它锈病群体过程中的分子表达特征分析。并运用BSMV-VIGS技术对其功能进行了初步研究。本研究的结果如下：
1.我们从小麦水源11与条锈菌CYR31的亲和cDNA文库中得到一条EST序列，其在Suwon11/CYR23的非亲和互作中显著的上调表达。利用电子克隆的方法，我们从水源11中扩增得到了TaAbc1基因。TaAbc1编码717个氨基酸的蛋白质，其等电点为9.38，分子量为80.35kDa。NCBI预测其具有Abc1-like蛋白家族的两个保守结构域，在羧基端有两个跨膜结构域和位于氨基端的一个叶绿体定位序列。序列分析表明，其与水稻和拟南芥聚成一个亚族。
2.不同器官中的基因表达模式显示该基因的表达是组织特异的，在茎中的表达最高，而在根中的表达最低只有叶中表达量的11%。在亲和与非亲和互作中研究该基因的表达模式。其中水源11分别与CYR31和CYR23组成亲和与非亲和组合，含有Lr47的美国春小麦与叶锈菌PBJL和秆锈菌QFCSC组成非亲和组合。TaAbc1在亲和组合和非亲和组合中的表达都上调并且在24小时达到了高峰。在条锈菌与小麦的非亲和互作中诱导其表达的倍数显著升高，说明该基因可能是条锈菌特异的HR中间产物。在外源乙烯和茉莉酸处理下基因的表达上调,并且伤诱导下有一个显著的上调，表明了其可能参与了伤诱导的信号通路。而在外源水杨酸和脱落酸作用下没有显著的变化，表明TaAbc1可能位于SA信号通路的上游或者独立于SA信号通路之外。
3.用100mM H2O2处理小麦植物，TaAbc1的转录水平有一个显著的升高。通过病毒诱导的基因沉默降低了水源11中的TaAbc1的表达，接条锈菌后细胞坏死的程度受到了抑制但并没有扭转整体的抗病性水平，表明该基因是包含在HR反应中，但整个寄主的整体抗病性不是HR依赖型的。这也表明了HR的信号通路和CYR23诱导的其它的防御反应通路是分支的。这些结果表明TaAbc1可能是一个在小麦防御真菌病原菌过程中的一个细胞死亡信号分子，并且TaAbc1介导的HR可能是锈病种类特异的。
抗秆锈基因Sr33从山羊草被渗透到普通小麦中而赋予了含有Sr33基因的小麦能抵御所有Ug99的衍生小种和全球不同地域的小麦秆锈菌小种。Lagudah从含有Sr33抗病基因的中国春中鉴定并克隆了Sr33基因。本研究通过BSMV-VIGS分析表明Sr33的抗病性功能是独立于RAR1, SGT1和HSP90的分子伴侣之外的。进而，沉默Sr33基因后提高了秆锈对小麦的侵染，这说明了Sr33在抗秆锈的防御途径中起主要作用。同时，沉默相邻的含有Exo70亚单位的基因后抗病反应型并没有发生变化，说明它在Sr33的抗病途径中是不需要的。
Plant hypersensitive response (HR), visualized as rapid death of host plant cells at theinfection sites, is a means of plant defense response to microbial pathogens. In plants, HR istriggered by incompatible interactions of host plants and corresponding pathogens. Stripe rust(Puccinia striiformis f. sp. tritici，Pst) can cause wheat stripe rust disease, and is aworldwide problem threatening wheat production. Therefore, searching for genes involved inwheat (Triticum aestivum L.) defense response to the stripe rust pathogen Pst is of greatlyimportance to elucidate the molecular mechanisms of HR and the signal pathway in theinteractions between the wheat and stripe rust.
To search for genes involved in wheat (Triticum aestivum L.) defense response to thestripe rust pathogen Pst, we identified and cloned a new wheat gene similar to the Abc1-likefamily. We present the results on molecular characteristics of this newly identified wheatAbc1-like gene, its expression profiles in response to Pst infection and abiotic elicitors and itsinvolvement in wheat defense response to a group of rust fungal pathogens. To test if TaAbc1has any functional role in wheat defense response to stripe rust, we knocked down theendogenous TaAbc1gene in Suwon11using the barley stripe mosaic virus induced genesilencing (BSMV-VIGS) assay.
1. From a cDNA library constructed from the wheat cultivar Suwon11infected with avirulent stripe rust pathotype CYR31，we selected an EST clone for further characterization inthis study due to its strong up-regulated expression in the incompatible interaction ofSuwon11/CYR23. The gene was successfully amplified from Suwon11after in silico. Thenew gene, designated as TaAbc1, encodes a717-amino acid,80.35kD protein. The TaAbc1protein contains two conserved domains shared by Abc1-like proteins, two trans-membranedomains at the C-terminal and a36-amino acid chloroplast targeting presequence at theN-terminal. Sequence analysis showed that TaAbc1were highly homologous to proteins inOryza sativa and Arabidopsis etc, and these proteins formed a subgroup.
2. Characterization of TaAbc1expression revealed that gene expression was tissue-specific. The highest TaAbc1transcript abundances were detected in stems. In contrast,roots had the lowest--only about11%of the level in leaves. To study the expression profilesof TaAbc1during the course of rust infections in both incompatible and compatibleinteractions, we challenged Suwon11with two Pst pathotypes, CYR23and CYR31. Inaddition, we included in the study two additional incompatible interactions with a USA springwheat genotype Scholar with a leaf rust resistance gene Lr47, inoculated with pathotype PBJLof the leaf rust pathogen (P. triticina) and pathotype QFCSC of the stem rust pathogen (P.graminis f. sp. tritici). TaAbc1was up-regulated and peaked at24hpi with rust challenge inboth incompatible and compatible interactions. High-fold induction was associated with thehypersensitive response (HR) triggered only by avirulent stripe rust pathotypes, suggestingTaAbc1is a rust-pathotype specific HR-mediator. An exogenous treatment of JA/ETup-regulated TaAbc1transcription but the increase was not as much as that detected in thedirectly wounded plants. The expression of TaAbc1was not affected by SA treatment,suggesting that TaAbc1was positioned either up-stream of SA or in an SA-independentmanner.
3. A significant increase in TaAbc1transcript level in the H2O2treated plant as early as at2hpt. Down-regulating TaAbc1expression by virus-induced gene silencing reduced necroticarea at infection sites but not the overall resistance level, suggesting TaAbc1was involved inHR against stripe rust, but overall host resistance is not HR-dependent. Our study suggestedthat signaling pathways to HR and other defense responses induced by CYR23in wheat arebranched. These observations invited the hypothesis that TaAbc1could be a commonly sharedcell-death signal used in wheat defense response to rust fungal pathogens. Our results tend tosuggest that TaAbc1-mediated HR was rust species and pathotype specific only in theincompatible interactions triggered by stripe rust.
Stem rust (Puccinia graminis f. sp. tritici) of wheat is a major threat to global foodsecurity and necessitates the continued development of new stem rust resistant varieties. Stemrust resistance gene Sr33derived from Aegilops tauschii and transferred into bread wheatconfers resistance against all stem rust pathogen derivatives of Ug99and diverse global wheatstem rust races. They identified the member Sr33responsible for stem rust resistance functionfrom the RGA Gene Cluster（Lagudah unpublished）.Characterization of Sr33-mediated stemrust resistance indicated that Sr33protein function is independent of RAR1, SGT1and HSP90chaperones by silencing genes encoding these chaperones in the hexaploid wheat line CSexpressing Sr33. Moreover, as the BSMV:Sr33treated plants displayed an increased susceptibility to stem rust infection, these data further validate the notion of this gene asproviding wheat with Sr33-dependent stem rust resistance. We also showed that silencing ofthe adjacent member carrying the exocyst70subunit did not compromise resistanceindicating the gene is not required for Sr33mediated resistance.
为了寻找小麦对条锈菌防御反应中的基因，我们克隆并鉴定了一个新的类似Abc1蛋白家族的基因。我们对小麦Abc1基因进行了条锈菌侵染的生物胁迫和非生物胁迫以及小麦防御其它锈病群体过程中的分子表达特征分析。并运用BSMV-VIGS技术对其功能进行了初步研究。本研究的结果如下：
1.我们从小麦水源11与条锈菌CYR31的亲和cDNA文库中得到一条EST序列，其在Suwon11/CYR23的非亲和互作中显著的上调表达。利用电子克隆的方法，我们从水源11中扩增得到了TaAbc1基因。TaAbc1编码717个氨基酸的蛋白质，其等电点为9.38，分子量为80.35kDa。NCBI预测其具有Abc1-like蛋白家族的两个保守结构域，在羧基端有两个跨膜结构域和位于氨基端的一个叶绿体定位序列。序列分析表明，其与水稻和拟南芥聚成一个亚族。
2.不同器官中的基因表达模式显示该基因的表达是组织特异的，在茎中的表达最高，而在根中的表达最低只有叶中表达量的11%。在亲和与非亲和互作中研究该基因的表达模式。其中水源11分别与CYR31和CYR23组成亲和与非亲和组合，含有Lr47的美国春小麦与叶锈菌PBJL和秆锈菌QFCSC组成非亲和组合。TaAbc1在亲和组合和非亲和组合中的表达都上调并且在24小时达到了高峰。在条锈菌与小麦的非亲和互作中诱导其表达的倍数显著升高，说明该基因可能是条锈菌特异的HR中间产物。在外源乙烯和茉莉酸处理下基因的表达上调,并且伤诱导下有一个显著的上调，表明了其可能参与了伤诱导的信号通路。而在外源水杨酸和脱落酸作用下没有显著的变化，表明TaAbc1可能位于SA信号通路的上游或者独立于SA信号通路之外。
3.用100mM H2O2处理小麦植物，TaAbc1的转录水平有一个显著的升高。通过病毒诱导的基因沉默降低了水源11中的TaAbc1的表达，接条锈菌后细胞坏死的程度受到了抑制但并没有扭转整体的抗病性水平，表明该基因是包含在HR反应中，但整个寄主的整体抗病性不是HR依赖型的。这也表明了HR的信号通路和CYR23诱导的其它的防御反应通路是分支的。这些结果表明TaAbc1可能是一个在小麦防御真菌病原菌过程中的一个细胞死亡信号分子，并且TaAbc1介导的HR可能是锈病种类特异的。
抗秆锈基因Sr33从山羊草被渗透到普通小麦中而赋予了含有Sr33基因的小麦能抵御所有Ug99的衍生小种和全球不同地域的小麦秆锈菌小种。Lagudah从含有Sr33抗病基因的中国春中鉴定并克隆了Sr33基因。本研究通过BSMV-VIGS分析表明Sr33的抗病性功能是独立于RAR1, SGT1和HSP90的分子伴侣之外的。进而，沉默Sr33基因后提高了秆锈对小麦的侵染，这说明了Sr33在抗秆锈的防御途径中起主要作用。同时，沉默相邻的含有Exo70亚单位的基因后抗病反应型并没有发生变化，说明它在Sr33的抗病途径中是不需要的。
Plant hypersensitive response (HR), visualized as rapid death of host plant cells at theinfection sites, is a means of plant defense response to microbial pathogens. In plants, HR istriggered by incompatible interactions of host plants and corresponding pathogens. Stripe rust(Puccinia striiformis f. sp. tritici，Pst) can cause wheat stripe rust disease, and is aworldwide problem threatening wheat production. Therefore, searching for genes involved inwheat (Triticum aestivum L.) defense response to the stripe rust pathogen Pst is of greatlyimportance to elucidate the molecular mechanisms of HR and the signal pathway in theinteractions between the wheat and stripe rust.
To search for genes involved in wheat (Triticum aestivum L.) defense response to thestripe rust pathogen Pst, we identified and cloned a new wheat gene similar to the Abc1-likefamily. We present the results on molecular characteristics of this newly identified wheatAbc1-like gene, its expression profiles in response to Pst infection and abiotic elicitors and itsinvolvement in wheat defense response to a group of rust fungal pathogens. To test if TaAbc1has any functional role in wheat defense response to stripe rust, we knocked down theendogenous TaAbc1gene in Suwon11using the barley stripe mosaic virus induced genesilencing (BSMV-VIGS) assay.
1. From a cDNA library constructed from the wheat cultivar Suwon11infected with avirulent stripe rust pathotype CYR31，we selected an EST clone for further characterization inthis study due to its strong up-regulated expression in the incompatible interaction ofSuwon11/CYR23. The gene was successfully amplified from Suwon11after in silico. Thenew gene, designated as TaAbc1, encodes a717-amino acid,80.35kD protein. The TaAbc1protein contains two conserved domains shared by Abc1-like proteins, two trans-membranedomains at the C-terminal and a36-amino acid chloroplast targeting presequence at theN-terminal. Sequence analysis showed that TaAbc1were highly homologous to proteins inOryza sativa and Arabidopsis etc, and these proteins formed a subgroup.
2. Characterization of TaAbc1expression revealed that gene expression was tissue-specific. The highest TaAbc1transcript abundances were detected in stems. In contrast,roots had the lowest--only about11%of the level in leaves. To study the expression profilesof TaAbc1during the course of rust infections in both incompatible and compatibleinteractions, we challenged Suwon11with two Pst pathotypes, CYR23and CYR31. Inaddition, we included in the study two additional incompatible interactions with a USA springwheat genotype Scholar with a leaf rust resistance gene Lr47, inoculated with pathotype PBJLof the leaf rust pathogen (P. triticina) and pathotype QFCSC of the stem rust pathogen (P.graminis f. sp. tritici). TaAbc1was up-regulated and peaked at24hpi with rust challenge inboth incompatible and compatible interactions. High-fold induction was associated with thehypersensitive response (HR) triggered only by avirulent stripe rust pathotypes, suggestingTaAbc1is a rust-pathotype specific HR-mediator. An exogenous treatment of JA/ETup-regulated TaAbc1transcription but the increase was not as much as that detected in thedirectly wounded plants. The expression of TaAbc1was not affected by SA treatment,suggesting that TaAbc1was positioned either up-stream of SA or in an SA-independentmanner.
3. A significant increase in TaAbc1transcript level in the H2O2treated plant as early as at2hpt. Down-regulating TaAbc1expression by virus-induced gene silencing reduced necroticarea at infection sites but not the overall resistance level, suggesting TaAbc1was involved inHR against stripe rust, but overall host resistance is not HR-dependent. Our study suggestedthat signaling pathways to HR and other defense responses induced by CYR23in wheat arebranched. These observations invited the hypothesis that TaAbc1could be a commonly sharedcell-death signal used in wheat defense response to rust fungal pathogens. Our results tend tosuggest that TaAbc1-mediated HR was rust species and pathotype specific only in theincompatible interactions triggered by stripe rust.
Stem rust (Puccinia graminis f. sp. tritici) of wheat is a major threat to global foodsecurity and necessitates the continued development of new stem rust resistant varieties. Stemrust resistance gene Sr33derived from Aegilops tauschii and transferred into bread wheatconfers resistance against all stem rust pathogen derivatives of Ug99and diverse global wheatstem rust races. They identified the member Sr33responsible for stem rust resistance functionfrom the RGA Gene Cluster（Lagudah unpublished）.Characterization of Sr33-mediated stemrust resistance indicated that Sr33protein function is independent of RAR1, SGT1and HSP90chaperones by silencing genes encoding these chaperones in the hexaploid wheat line CSexpressing Sr33. Moreover, as the BSMV:Sr33treated plants displayed an increased susceptibility to stem rust infection, these data further validate the notion of this gene asproviding wheat with Sr33-dependent stem rust resistance. We also showed that silencing ofthe adjacent member carrying the exocyst70subunit did not compromise resistanceindicating the gene is not required for Sr33mediated resistance.
引文
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