摘要
禾谷镰刀菌引起的赤霉病是小麦的重要病害之一,严重地影响小麦的产量及品质。因此,对抗小麦赤霉病的基因以及赤霉病防卫相关基因的克隆和鉴定,不仅可以增加我们对赤霉病抗性机制的理解,还可以丰富小麦抗赤霉病育种改良的候选基因。
为了挖掘抗病相关的基因,我们对响应生物以及非生物胁迫的Jacalin相关的凝集素基因(JRL)进行了鉴定。通过对NCBI中小麦EST数据库的检索,共得到53个编码JRL的EST重叠群和singleton。根据JRL蛋白的结构域的数目,我们把水稻、小麦和拟南芥的JRL基因划分成为四组。JRL基因的外显子和内含子结构多样化程度很高。尽管JRL蛋白的氨基酸序列一致性低且蛋白结构域构架变异大,但其Jacalin结构域具有较高的保守性。根据对水稻和拟南芥的芯片数据和MPSS数据的分析以及小麦JRL基因的定量表达分析,我们发现JRL基因的表达具有时空特异性,这类基因可能在植物生长和发育的特定时期发挥功能。不同物种的JRL基因家族成员几乎都能响应多种生物、非生物胁迫以及植物激素的刺激；在四组JRL基因中,随着JRL蛋白中Jacalin结构域数目的增多,每组中响应不同种类胁迫的JRL基因的比例逐渐增多或减少。在水稻和拟南芥中,大多数胁迫响应的JRL基因的启动子都包含相应的胁迫响应顺式作用元件。不同的JRL可能具有一定的功能冗余,同时也进化了自己独特的功能。
为了了解JRL基因与植物抗病性间的关系,我们从抗赤霉病品种望水白中克隆到一个表达受赤霉菌诱导的JRL基因(TaJRL1)。该基因编码一个包含两个Jacalin结构域的新酸性多肽。亚细胞定位结果表明它主要定位于细胞核中。TaJRL1响应病原菌侵染、植物激素处理以及干旱和热激等非生物胁迫。当应用水杨酸(SA)生物合成抑制剂paclobutrazol (PAC),或茉莉酸(JA)生物合成抑制剂diethyldithiocarbamic acid (DIECA)时,TaJRL1的表达受到明显抑制。应用BSMV病毒介导的基因沉默系统(VIGS)降低内源TaJRL1基因的表达时,小麦植株不仅更感死体寄生真菌赤霉菌,也更感活体寄生真菌白粉菌。转TaJRL1的拟南芥植株中游离SA和JA的含量增高,同时也增强了对赤霉菌和灰霉菌的抗性。依赖SA和JA防卫途径中的基因的抑制或上调分别与这些植株的感病性或抗病性明显相关。TaJRL1有可能调控了JA和SA的生物合成,进而调控SA和JA依赖的防卫途径的协同作用,从而改变植物的基础抗病反应。
与JRL蛋白类似,钙网联蛋白(calreticulin, CRT)是植物凝集素中的另一个亚家族。我们从望水白中克隆得到三个CRT基因(TaCRT1-3)。系统进化分析表明CRT在单子叶和双子叶植物中具有较好的保守性,并可划分为CRT1/2和CRT3两个亚家族。TaCRT在生殖组织中的表达丰度很高,TaCRT1和TaCRT3在营养生长组织中也具有较高的表达丰度。赤霉菌侵染和赤霉菌毒素DON处理能诱导TaCRT的表达。TaCRT1响应JA的刺激,TaCRT2响应JA和ETH的刺激,TaCRT3响应SA和JA的刺激。在拟南芥中过量表达TaCRTl降低了植株对灰霉病的抗性,当转入去除内质网定位信号和滞留信号的TaCRT1(dsTaCRTl)后植株对灰霉菌的抗性增强。免疫胶体金定位表明TaCRT1蛋白定位于内质网和细胞质中。表达结果显示,在转dsTaCRTl的拟南芥植株中,钙调素连接蛋白基因CBP60g、 SA依赖的防卫途径中的PAD4.ICS1、EDS5、PR1和PR2的表达都明显上调,而TaCRTl的过量表达下调了这些基因的表达。我们推测TaCRTl可能通过对CBP60g的调控进而影响水杨酸依赖的防卫信号途径。
Scab disease, caused by Fusarium graminearum Schwabe, is a destructive disease of wheat worldwide, result in great yield losses and poor grain quality. So, isolation and characterization of the defense-related genes for scab would not only increase our understanding of the scab resistance mechanisms, but also enrich the candidate genes for fighting against the disastrous scab disease.
To disentomb defense-related genes, we identified the jacalin-related lectin (JRL) gene response to various biotic and/or abiotic stresses. We identified53EST contigs and sigletons encoding JRL genes in wheat through mining the NCBI wheat EST database. Based on number of jacalin domain of wheat, rice and Arabidopsis, JRL genes can be classified into four groups. These JRL genes have diverse exon-intron structures. Although the overall pairwise protein sequence identity is modest and the overall architecture of JRLs proteins showed marked individual differences, the jacalin domain display highly conserved peculiarities. Microarray and MPSS data and real-time PCR revealed that JRL genes showed different tempo-spatial expression, suggesting that these genes may function at different stages of plant growth and development. Meanwhile, many JRL genes were response to various biotic and/or abiotic stress conditions and phytohormone applied, and the ratio of stress response JRL genes in every group altered gradually with the increasing of number of jacalin domain. Major stress-responsive OsJRL and At JRL genes contained putative corresponding stress-responsive cis-elements. These results showed that JRL genes had redundant but distinct functions.
To research the disease resistance of JRL genes, we had identified a Triticum-specific jacalin-related lectin(TaJRL1) gene from scab-resistant wheat (Triticum aestivum L) cv. Wangshuibai. TaJRLl encodes a novel acidic polypeptide with two jacalin domains, and it was located mainly in the nuclei. Pathogen infection, phytohormone treatments, drought and heat shock stresses induced expression of TaJRLl; while application of the salicylic acid (SA) biosynthesis inhibitor paclobutrazol (PAC), or jasmonic acid (JA) biosynthesis inhibitor diethyldithiocarbamic acid (DIECA) substantially inhibited its expression. Attenuating TaJRLl through virus-induced gene silencing (VIGS) enhanced susceptibility to facultative fungal pathogen Fusarium graminearum and biotrophic fungal pathogen Blumeria graminis. Arabidopsis plants transformed with TaJRLl displayed increased resistance to Fusarium graminearum and Botrytis cinerea. The JA and SA levels in these transgenic Arabidopsis thaliana plants rose significantly. The loss or increase of disease resistances because of alternation of TaJRL1function correlated well with attenuation or enhancement of the SA-and JA-dependent defense signaling pathways. These results suggested that TaJRLl appears to have a role in feedback amplification of the defense signaling pathways.
Similaring to JRL, calreticulin (CRT) is also a kind of plant lectins. In this study, we characterized three CRT gene family memers from wheat cv. Wangshuibai. Phylogenetic analysis showed that CRT proteins were well conserved among monocots and dicots, and were grouped into CRT1/2and CRT3subfamilies. Their transcripts were mainly detected in reproductive tissues, and TaCRTl/3were also highly expressed in vegetative tissues. Fusarium grminearum infection and elicitor DON treatment induced TaCRT genes expression. TaCRT1was up-regulated by JA treatment, TaCRT2was induced by JA and ETH treatments, TaCRT3was up-regulated by SA and JA treatment. Overexpression of TaCRTl in Arabidopsis decreased resistance to Botrytis cinerea, however, dsTaCRTl transgenic Arabidopsis thaliana enhanced resistance. Immunogold localization showed that TaCRT1and dsTaCRTl targeted to ER and cytoplasmic, and cytoplasmic, respectively. qRT-PCR showed that calcium-binding protein gene (CBP60g) and genes (PAD4, ICS1, EDS5, PR1and PR2) in SA-dependent defense signaling pathway were up-regulated in dsTaCRTl transgenic plants. These genes were down-regulated in TaCRTl transgenic plants. Our results indicated that TaCRT1might regulate SA-mediated defense signaling pathway by CBP60g gene.
引文
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