WRKY转录因子SIDRW1和NAC转录因子SISRN1在番茄抗性反应中的功能研究
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摘要
转录因子作为复杂的信号网络中的成员,在调节植物应对来自周围环境因子胁迫的过程中发挥着重要的作用。其中,WRKY、NAC等转录因子家族成员在植物生长发育和抗逆等过程中发挥着重要的调节功能。而目前关于这两个转录因子家族的研究主要集中在水稻和拟南芥中,番茄等其他植物中的研究很少。本文从番茄中克隆了12个NAC和6个WRKY转录因子基因,并且主要利用病毒诱导的基因沉默(VIGS)等技术鉴定了它们在番茄抵抗病原菌以及非生物胁迫过程中的作用,初步筛选出2个具有重要调节功能的基因SIDRW1(defense-related WRKY1)和SISRN1(stress-related NAC1)。
实验结果表明,WRKY转录因子SlDRW1在番茄与灰霉互作过程中是一个正调控因子。分析发现,、VRKY转录因子基因SlDRW1编码一个由360个氨基酸组成的蛋白,根据推测其分子量大约为39.7kDa,等电点为8.13。而且,它在序列上与另外两个番茄WRKY转录因子基因SlWRKY1和SlWRKY2分别具有99%和91%的相似度。进一步对SlDRW1和SlWRKY1的ORF氨基酸序列比对分析发现,这两个基因为同一基因。通过实验手段验证了SlDRW1的转录激活活性,结果表明,它可能是一个转录抑制子。GFP融合蛋白分析显示,该基因定位于细胞核内。接种灰霉病菌后,SVDRW1在番茄叶片内的表达在接种后24h (24hpi)突然升高,48hpi达到高峰,从24hpi开始持续保持在较高表达水平；然而,接种Pst DC3000后仅在48hpi略有上调表达,其余时间点均恢复到正常表达水平。用SA、JA诱导后均有不同水平的上调表达但ACC处理后,SlDRW1的表达水平基本上没有明显变化。为了进一步验证SlDRW1基因在番茄与病原菌互作中的功能我们构建了其VIGS表达载体。Realtime-PCR实验表明,利用VIGS技术,可以使SlDRW1在番茄中的转录水平平均下调50-80%,最高可达90%以上。SlDRW1沉默的番茄叶片离体接种灰霉后,病斑直径明显比对照大。我们同时构建该基因瞬时过表达载体,结果,瞬间过表达处理过的叶片抵抗灰霉的能力明显提高,主要表现在病斑较对照小。SlDRW1沉默的番茄植株接种灰霉48h后,体内活性氧积累明显比对照少。为了进一步探明SlDRW1发挥作用的内在机制,验证了接种灰霉菌以后,几个PR基因在处理和对照植物中的表达差异。其中,PR1a、PR1b、PR2a、PR3a、PR5、PR7均上调表达,而PR2b、PR3b的表达与对照没有差异。这表明SlDRW1可能通过调节活性氧和部分PR基因的表达而影响植物对病原物的抗性。虽然,SlDRW1在番茄与灰霉(Botrytis cinerea)互作中是一个正调控因子,但对番茄抵御细菌性叶斑病、干旱和氧化胁迫中没有明显作用。
同时发现,NAC转录因子SlSRN1在番茄与灰霉和细菌性叶斑病互作中均为正调控因子,而且,它在番茄抵抗干旱胁迫过程中是一个负调控因子。该基因在灰霉菌、SA和JA诱导下均有一定程度的上调表达,而Pst DC3000和ACC诱导后其表达与对照相比较却没有明显差异。通过转录激活活性分析进一步验证了其转录因子身份。亚细胞定位实验表明,SlSRN1定位于细胞核。实验结果表明VIGS沉默番茄植株对灰霉和细菌性叶斑病菌抵抗力均下降。这说明S(?)lSRN1在番茄抵抗这两种病原菌过程中发挥着重要作用。灰霉接种后SlSRN1基因沉默的番茄植株体内PR5和PR7表达量明显上调,说明SlSRN1可能通过调节病程相关蛋白的表达而在番茄抗病过程中发挥作用。
Complex signaling networks that involve transcription factors play a central role in plant responses to surrounding crisises. The members of several transcription factor families, such as ethylene-responsive-element-binding factors (ERF)、basic-domain leucine-zipper (bZIP)、MYB、NAC and WRKY proteins have proven to involve in the plant defense response against abiotic and biotic stresses. Among them, NAC and WRKY are two of the biggest and most intensely researched transcription factor families in plants. However, most of the studies had been focused on rice and Arabidopsis, researches in these fields on tomato was rare. In this paper,12NAC and6WRKY transcription factors were cloned, and their VIGS vectors were constructed. Taken the VIGS as main tool, we identified a WRKY transcription factors gene, SlDRW1(defense-related WRKY1) and a NAC transcription factors gene SlSRN1(stress-related NAC1), respectively, which involved in tomato-disease interactions.
The WRKY transcription factor gene SlDRW1is1731bp in size with a predicted open reading frame of1083bp. SlDRW1encodes a protein of360amino acids and the deduced protein contains all conserved domains characteristics of the WRKY group Ⅱ members. And the amino acids BLAST results demonstrated that SlDRW1has a99%and91%similarity in sequence with tomato other two WRKY transcription factors SlWRKY1and SlWRKY2respectively. Further alignment analysis of amino acid sequences for the ORF of SlDRW1and SlWRKY1, showed they are identical. Transactivation analysis demonstrated that SlDRW1can not activate reporter gene expression in yeast, indicating that SlDRW1maybe a transcriptional repressor. Subcellular localization analysis using GFP fusion protein showed that the SlDRW1protein is localized in the nucleus of Nicotiana benthamiana and tomato epidermal cells.
The expression of SlDRW1gene was up-regulated dramatically24hours post infection (DPI) with Botrytis cinerea and peaked at48DPI. It also increased slightly by treatment with Pseudomonas syringae pv. tomato (Pst) DC3000, salicylic acid (SA) and jasmonate (JA), but not by1-aminocyclopropane-1-carboxylic acid (ACC), suggesting that SlDRW1may be involved in regulation of defense response in tomato.
Then VIGS was adopted as vehicle to reveal the roles of SlDRW1in tomato-disease interaction. Quantitative real-time PCR analysis demonstrated that the transcripts of SlDRW1could be reduced by50-80%,even more than90%,around4weeks post VIGS treatment. After inoculation with Botrytis cinerea, the SlDRW1VIGS silenced tomatoes showed more susceptible phenotype than the control plants with bigger disease size. So we drew a conclusion that SlDRW1confers resistance to tomato against Botrytis cinerea. And the transient over-expression study confirmed the result. To grope for the mechanism underlying the phenomenon, the expression of active oxygen and pathogenesis-related proteins (PR) were tested. SlDRW1may modulate tomato defense response against Botrytis cinerea by regulation of the PR genes and active oxygen species expression.Nevertheless, SlDRW1has no effect on the response of tomato against Pst DC3000, drought and oxidative stress.
The NAC transcription factor SlSRN1which contains a1470bp ORF and encodes a protein of490AA was also cloned and identified. The NAC gene endow resistance to tomato against necrotrophic fungus Botrytis cinerea and bacteria Pst DC3000, however, it act as a negative regulator in tomato drought responses. The expression of SlSRN1increased upon Botrytis cinerea and Pst DC3000infection, and the exogenous application of JA,SA, but not ACC. SlSRN1is a transcriptional activator since the transactivation analysis showed that SlSRN1activated reporter gene expression in yeast. Subcellular localization analysis showed that the SlSRN1protein is localized in the nucleus of cells. The mRNA products of SlSRN1in tomatoes can be reduced by an average of50%with VIGS technique. When the VIGS silenced tomatoes inoculated with B. cinerea, they showed more susceptible phenotypes and higher pathogen growth rate, indicating SlSRN1confers resistance to tomatoes against this pest. As to Pst DC3000, the silenced plants displayed similar symptoms, however, the bacteria grow much faster on SlSRN1silenced tomatoes, which made us draw a conclusion that SlSRN1acts as a positive regulator in tomato-Pst DC3000interaction. After withholding water for5days, SlSRN1silenced tomatoes showed less susceptible symptoms, indicating SlSRN1is a negative regulator in tomato response to drought. Results also hint that SlSRN1may be involved in regulating the expression of PR genes, via which to give rise to resistance of tomatoes against pathogens.
实验结果表明,WRKY转录因子SlDRW1在番茄与灰霉互作过程中是一个正调控因子。分析发现,、VRKY转录因子基因SlDRW1编码一个由360个氨基酸组成的蛋白,根据推测其分子量大约为39.7kDa,等电点为8.13。而且,它在序列上与另外两个番茄WRKY转录因子基因SlWRKY1和SlWRKY2分别具有99%和91%的相似度。进一步对SlDRW1和SlWRKY1的ORF氨基酸序列比对分析发现,这两个基因为同一基因。通过实验手段验证了SlDRW1的转录激活活性,结果表明,它可能是一个转录抑制子。GFP融合蛋白分析显示,该基因定位于细胞核内。接种灰霉病菌后,SVDRW1在番茄叶片内的表达在接种后24h (24hpi)突然升高,48hpi达到高峰,从24hpi开始持续保持在较高表达水平；然而,接种Pst DC3000后仅在48hpi略有上调表达,其余时间点均恢复到正常表达水平。用SA、JA诱导后均有不同水平的上调表达但ACC处理后,SlDRW1的表达水平基本上没有明显变化。为了进一步验证SlDRW1基因在番茄与病原菌互作中的功能我们构建了其VIGS表达载体。Realtime-PCR实验表明,利用VIGS技术,可以使SlDRW1在番茄中的转录水平平均下调50-80%,最高可达90%以上。SlDRW1沉默的番茄叶片离体接种灰霉后,病斑直径明显比对照大。我们同时构建该基因瞬时过表达载体,结果,瞬间过表达处理过的叶片抵抗灰霉的能力明显提高,主要表现在病斑较对照小。SlDRW1沉默的番茄植株接种灰霉48h后,体内活性氧积累明显比对照少。为了进一步探明SlDRW1发挥作用的内在机制,验证了接种灰霉菌以后,几个PR基因在处理和对照植物中的表达差异。其中,PR1a、PR1b、PR2a、PR3a、PR5、PR7均上调表达,而PR2b、PR3b的表达与对照没有差异。这表明SlDRW1可能通过调节活性氧和部分PR基因的表达而影响植物对病原物的抗性。虽然,SlDRW1在番茄与灰霉(Botrytis cinerea)互作中是一个正调控因子,但对番茄抵御细菌性叶斑病、干旱和氧化胁迫中没有明显作用。
同时发现,NAC转录因子SlSRN1在番茄与灰霉和细菌性叶斑病互作中均为正调控因子,而且,它在番茄抵抗干旱胁迫过程中是一个负调控因子。该基因在灰霉菌、SA和JA诱导下均有一定程度的上调表达,而Pst DC3000和ACC诱导后其表达与对照相比较却没有明显差异。通过转录激活活性分析进一步验证了其转录因子身份。亚细胞定位实验表明,SlSRN1定位于细胞核。实验结果表明VIGS沉默番茄植株对灰霉和细菌性叶斑病菌抵抗力均下降。这说明S(?)lSRN1在番茄抵抗这两种病原菌过程中发挥着重要作用。灰霉接种后SlSRN1基因沉默的番茄植株体内PR5和PR7表达量明显上调,说明SlSRN1可能通过调节病程相关蛋白的表达而在番茄抗病过程中发挥作用。
Complex signaling networks that involve transcription factors play a central role in plant responses to surrounding crisises. The members of several transcription factor families, such as ethylene-responsive-element-binding factors (ERF)、basic-domain leucine-zipper (bZIP)、MYB、NAC and WRKY proteins have proven to involve in the plant defense response against abiotic and biotic stresses. Among them, NAC and WRKY are two of the biggest and most intensely researched transcription factor families in plants. However, most of the studies had been focused on rice and Arabidopsis, researches in these fields on tomato was rare. In this paper,12NAC and6WRKY transcription factors were cloned, and their VIGS vectors were constructed. Taken the VIGS as main tool, we identified a WRKY transcription factors gene, SlDRW1(defense-related WRKY1) and a NAC transcription factors gene SlSRN1(stress-related NAC1), respectively, which involved in tomato-disease interactions.
The WRKY transcription factor gene SlDRW1is1731bp in size with a predicted open reading frame of1083bp. SlDRW1encodes a protein of360amino acids and the deduced protein contains all conserved domains characteristics of the WRKY group Ⅱ members. And the amino acids BLAST results demonstrated that SlDRW1has a99%and91%similarity in sequence with tomato other two WRKY transcription factors SlWRKY1and SlWRKY2respectively. Further alignment analysis of amino acid sequences for the ORF of SlDRW1and SlWRKY1, showed they are identical. Transactivation analysis demonstrated that SlDRW1can not activate reporter gene expression in yeast, indicating that SlDRW1maybe a transcriptional repressor. Subcellular localization analysis using GFP fusion protein showed that the SlDRW1protein is localized in the nucleus of Nicotiana benthamiana and tomato epidermal cells.
The expression of SlDRW1gene was up-regulated dramatically24hours post infection (DPI) with Botrytis cinerea and peaked at48DPI. It also increased slightly by treatment with Pseudomonas syringae pv. tomato (Pst) DC3000, salicylic acid (SA) and jasmonate (JA), but not by1-aminocyclopropane-1-carboxylic acid (ACC), suggesting that SlDRW1may be involved in regulation of defense response in tomato.
Then VIGS was adopted as vehicle to reveal the roles of SlDRW1in tomato-disease interaction. Quantitative real-time PCR analysis demonstrated that the transcripts of SlDRW1could be reduced by50-80%,even more than90%,around4weeks post VIGS treatment. After inoculation with Botrytis cinerea, the SlDRW1VIGS silenced tomatoes showed more susceptible phenotype than the control plants with bigger disease size. So we drew a conclusion that SlDRW1confers resistance to tomato against Botrytis cinerea. And the transient over-expression study confirmed the result. To grope for the mechanism underlying the phenomenon, the expression of active oxygen and pathogenesis-related proteins (PR) were tested. SlDRW1may modulate tomato defense response against Botrytis cinerea by regulation of the PR genes and active oxygen species expression.Nevertheless, SlDRW1has no effect on the response of tomato against Pst DC3000, drought and oxidative stress.
The NAC transcription factor SlSRN1which contains a1470bp ORF and encodes a protein of490AA was also cloned and identified. The NAC gene endow resistance to tomato against necrotrophic fungus Botrytis cinerea and bacteria Pst DC3000, however, it act as a negative regulator in tomato drought responses. The expression of SlSRN1increased upon Botrytis cinerea and Pst DC3000infection, and the exogenous application of JA,SA, but not ACC. SlSRN1is a transcriptional activator since the transactivation analysis showed that SlSRN1activated reporter gene expression in yeast. Subcellular localization analysis showed that the SlSRN1protein is localized in the nucleus of cells. The mRNA products of SlSRN1in tomatoes can be reduced by an average of50%with VIGS technique. When the VIGS silenced tomatoes inoculated with B. cinerea, they showed more susceptible phenotypes and higher pathogen growth rate, indicating SlSRN1confers resistance to tomatoes against this pest. As to Pst DC3000, the silenced plants displayed similar symptoms, however, the bacteria grow much faster on SlSRN1silenced tomatoes, which made us draw a conclusion that SlSRN1acts as a positive regulator in tomato-Pst DC3000interaction. After withholding water for5days, SlSRN1silenced tomatoes showed less susceptible symptoms, indicating SlSRN1is a negative regulator in tomato response to drought. Results also hint that SlSRN1may be involved in regulating the expression of PR genes, via which to give rise to resistance of tomatoes against pathogens.
引文
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