陆地棉中NAC家族与叶片衰老和胁迫应答相关功能的遗传学分析
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摘要
NAC (NAM, ATAF,以及CUC)是植物特有的一类转录因子,NAC转录因子不仅参与植物生长发育的调控,而且在植物抗逆反应中具有重要的调控作用。NAC家族蛋白在各种植物发育过程中也起到一定作用,如叶片衰老,花的形态,逆境诱导开花,茎端分生组织的发育,茎分枝,侧根发育,激素信号等。NAC蛋白结构域与植物响应生物和非生物胁迫息息相关,包括盐分,低温,干旱,机械损伤以及病毒感染等。同时,NAC也参与纤维发育以及木质部的形成。尽管在模式植物中(拟南芥和水稻)鉴定了一些NAC转录因子的功能,但是在棉花中,特别是叶片衰老方面,NAC的功能尚未见报导。在本实验中,我们鉴定了陆地棉中NAC基因家族,并探索了其在棉花叶片衰老以及各种抗逆反应中的功能拟南芥中过表达GhNAC8和GhNAC9在叶片衰老时表现出重要作用。利用荧光定量PCR实验分析表明：这些基因在叶片衰老过程中以及抗逆应答中高水平表达。通过结合荧光定量PCR实验,免疫印迹分析以及拟南芥和烟草的过表达实验,NAC家族中的成员之一GhNAC7在叶片衰老中有重要作用。GhNAC的启动子有许多响应生理和环境因素诱导的顺式作用元件。
本文的主要研究结果总结如下：
1. GhNAC8-GhNAC17在叶片衰老以及应对胁迫反应过程中的作用
我们从陆地棉中分离得到了GhNAC8-GhNAC17基因,并考察了其在叶片衰老过程以及胁迫应答中的功能。这些GhNAC基因预测得到的氨基酸序列与其他物种NAC家族序列具有很高的相似性,基于这些特征,将GhNAC基因家族分成三个亚类。这些GhNAC基因中除了GhNAC10以及GhNAC13,都在衰老叶片中有很高的表达水平,而GhNAC10以及GhNAC13在棉花开花后(DPA)25天的植物纤维中具有很高的表达水平。与野生型相比,在拟南芥中异位表达GhNAC8和GhNAC9导致叶片提前衰老,这结果表明其在拟南芥的叶片衰老过程中具有生物学功能。这十个GhNAC基因在自然和诱导的叶片衰老过程中表现出不同的表达模式和表达水平。通过定量PCR和启动子分析表明,脱落酸,乙烯,干旱,盐分,低温,高温以及其他激素处理都能诱导这些基因的表达。
2.棉花NAC基因家族(GhNAC18-77)在多种非生物胁迫过程中的表达模式分析从陆地棉中一共分离得到了60个全长的GhNAC基因,通过系统发育分析,他们可以被分为七个不同的亚类。我们还构建了GhNAC基因的系统发育树,不同的亚类中,他们的结构域之间具有相似性。这些基因位于D亚类基因组的第十三条染色体上。通过荧光定量PCR分析这些基因的组织特异性,结果显示：大部分NAC基因的表达模式具有特定的时空特异性。另外,本实验还通过荧光定量PCR分析了这些GhNAC基因在叶片衰老和乙烯,脱落酸,赤霉素,干旱以及盐分处理后的表达变化。
3.一个新的GhNAC家族转录因子,GhNAC7基因在叶片衰老过程中的功能。
我们分析了编码NAC家族转录因子的GhNAC7基因的功能。通过荧光定量PCR和免疫印迹分析基因的组织特异性,结果显示：该基因与棉花叶片衰老过程具有密切关系。同时该基因也受乙烯、脱落酸、干旱、盐分、低温以及高温的诱导表达。在模式植物拟南芥以及烟草中过表达该基因,转基因植株比野生型植株表现出明显的衰老症状。利用免疫印迹实验,在大肠杆菌BL21中诱导表达GhNAC7蛋白,表明蛋白分子量大概为29kDa。以上结果表明GhNAC7在棉花叶片衰老中发挥了重要作用。
NAC (NAM, ATAF, and CUC) is a plant-specific transcription factor family with diverse roles in plant development and stress regulation. NAC family proteins also contribute to a variety of developmental processes in plants, such as leaf senescence, floral morphogenesis, stress induced flowering, development of shoot apical meristem, shoot branching determination, lateral root development, and hormone signaling. The NAC domain proteins has been associated with plant responses to biotic and abiotic stresses including salinity, cold shock, drought, mechanical wounding, and viral infections. Some other areas where their involvement has been reported are fiber development, xylogenesis and wood formation. Despite a few NAC TFs in Arabidopsis and Oryza model plants have been characterized for their functions, no reports are available for their functional characterization in cotton specifically for leaf senescence.
In this study, we identified the GhNAC gene family, characterized for leaf senescence and diverse stresses in cotton(Gossypium hirsutum L.). The qRT-PCR analysis revealed that these genes have high expression during leaf senescence and response to stresses. GhNAC8and GhNAC7overexpression lines in Arabidopsis depict essential role during leaf senescence. GhNAC7being a member of cotton NAC family has important functions in leaf senescence as elucidated by qRT-PCR, immunoblot analysis, overexpression in Arabidopsis and tobacco. GhNAC genes promoter regions possess several cis-elements inducible by physiological and environmental factors.
The main results are summarized as follows:
1. Characterization of GhNAC8-GhNACl7genes for leaf senescence and responses to stresses
Stress-responsive NAC genes (GhNAC8-GhNAC17) isolated from cotton (Gossypium hirsutum L.) were characterised in the context of leaf senescence and stress tolerance. Based on the sequence characterisation, these GhNACs could be classified into three groups belonging to three known NAC sub-families. Their predicted amino acid sequences exhibited similarities to NAC genes from other plant species. All GhNAC genes were highly expressed in senescent leaves, except GhNAC10and GhNAC13, which had higher expression levels in fibres collected from25days post anthesis (DPA) plants. Ectopic expression of GhNAC8and GhNAC9show early leaf senescence compared to wild-types plant, revealed biological function for leaf senescence in Arabidopsis. The ten GhNAC genes displayed differential expression patterns and levels during natural and induced leaf senescence. Quantitative RT-PCR and promoter analyses suggest that these genes are induced by ABA, ethylene, drought, salinity, cold, heat and other hormonal treatments.
2. Expression analysis of NAC family gene (GhNAC18-77) for multiple abiotic stresses in cotton
A total of60full-length GhNAC genes were isolated from upland cotton, and they were phylogeneticly clustered into seven distinct subfamilies. We also constructed the phylogenetic tree for GhNAC genes and their motif, which show similarities among the subfamilies. The60GhNAC genes were located in the chromosome of D sub-genome and were found to be distributed on13chromosomes. Majority of the NACs showed specific temporal and spatial expression patterns for tissue-specific studies based on qRT-PCR analyses. Furthermore, the roles of GhNAC genes were monitored by qRT-PCR for leaf senescence, ethylene, ABA, GA3, drought and salinity.
3. GhNAC7, a novel GhNAC family transcription factor gene functions in leaf senescence
GhNAC7, a gene encoding a NAC family transcription factor was studied for functional analysis. Expression of this gene is closely associated with the leaf senescence process of cotton as revealed by tissue-specific and natural leaf senescence studies through qRT-PCR and immunoblot assays. The GhNAC7expression also induced by ethylene, ABA, drought, salinity, cold and high temperature treatments in cotton. In two model plant species, Arabidopsis and tobacco overexpression of GhNAC7showed visible symptoms for leaf senescence. Protein expression in BL21E. Coli and immunoblot assay in senescent leaf confirmed that molecular weight of GhNAC7protein was approximately29kDa. Our data strongly suggest that GhNAC7play an important role in cotton leaf senescence.
本文的主要研究结果总结如下：
1. GhNAC8-GhNAC17在叶片衰老以及应对胁迫反应过程中的作用
我们从陆地棉中分离得到了GhNAC8-GhNAC17基因,并考察了其在叶片衰老过程以及胁迫应答中的功能。这些GhNAC基因预测得到的氨基酸序列与其他物种NAC家族序列具有很高的相似性,基于这些特征,将GhNAC基因家族分成三个亚类。这些GhNAC基因中除了GhNAC10以及GhNAC13,都在衰老叶片中有很高的表达水平,而GhNAC10以及GhNAC13在棉花开花后(DPA)25天的植物纤维中具有很高的表达水平。与野生型相比,在拟南芥中异位表达GhNAC8和GhNAC9导致叶片提前衰老,这结果表明其在拟南芥的叶片衰老过程中具有生物学功能。这十个GhNAC基因在自然和诱导的叶片衰老过程中表现出不同的表达模式和表达水平。通过定量PCR和启动子分析表明,脱落酸,乙烯,干旱,盐分,低温,高温以及其他激素处理都能诱导这些基因的表达。
2.棉花NAC基因家族(GhNAC18-77)在多种非生物胁迫过程中的表达模式分析从陆地棉中一共分离得到了60个全长的GhNAC基因,通过系统发育分析,他们可以被分为七个不同的亚类。我们还构建了GhNAC基因的系统发育树,不同的亚类中,他们的结构域之间具有相似性。这些基因位于D亚类基因组的第十三条染色体上。通过荧光定量PCR分析这些基因的组织特异性,结果显示：大部分NAC基因的表达模式具有特定的时空特异性。另外,本实验还通过荧光定量PCR分析了这些GhNAC基因在叶片衰老和乙烯,脱落酸,赤霉素,干旱以及盐分处理后的表达变化。
3.一个新的GhNAC家族转录因子,GhNAC7基因在叶片衰老过程中的功能。
我们分析了编码NAC家族转录因子的GhNAC7基因的功能。通过荧光定量PCR和免疫印迹分析基因的组织特异性,结果显示：该基因与棉花叶片衰老过程具有密切关系。同时该基因也受乙烯、脱落酸、干旱、盐分、低温以及高温的诱导表达。在模式植物拟南芥以及烟草中过表达该基因,转基因植株比野生型植株表现出明显的衰老症状。利用免疫印迹实验,在大肠杆菌BL21中诱导表达GhNAC7蛋白,表明蛋白分子量大概为29kDa。以上结果表明GhNAC7在棉花叶片衰老中发挥了重要作用。
NAC (NAM, ATAF, and CUC) is a plant-specific transcription factor family with diverse roles in plant development and stress regulation. NAC family proteins also contribute to a variety of developmental processes in plants, such as leaf senescence, floral morphogenesis, stress induced flowering, development of shoot apical meristem, shoot branching determination, lateral root development, and hormone signaling. The NAC domain proteins has been associated with plant responses to biotic and abiotic stresses including salinity, cold shock, drought, mechanical wounding, and viral infections. Some other areas where their involvement has been reported are fiber development, xylogenesis and wood formation. Despite a few NAC TFs in Arabidopsis and Oryza model plants have been characterized for their functions, no reports are available for their functional characterization in cotton specifically for leaf senescence.
In this study, we identified the GhNAC gene family, characterized for leaf senescence and diverse stresses in cotton(Gossypium hirsutum L.). The qRT-PCR analysis revealed that these genes have high expression during leaf senescence and response to stresses. GhNAC8and GhNAC7overexpression lines in Arabidopsis depict essential role during leaf senescence. GhNAC7being a member of cotton NAC family has important functions in leaf senescence as elucidated by qRT-PCR, immunoblot analysis, overexpression in Arabidopsis and tobacco. GhNAC genes promoter regions possess several cis-elements inducible by physiological and environmental factors.
The main results are summarized as follows:
1. Characterization of GhNAC8-GhNACl7genes for leaf senescence and responses to stresses
Stress-responsive NAC genes (GhNAC8-GhNAC17) isolated from cotton (Gossypium hirsutum L.) were characterised in the context of leaf senescence and stress tolerance. Based on the sequence characterisation, these GhNACs could be classified into three groups belonging to three known NAC sub-families. Their predicted amino acid sequences exhibited similarities to NAC genes from other plant species. All GhNAC genes were highly expressed in senescent leaves, except GhNAC10and GhNAC13, which had higher expression levels in fibres collected from25days post anthesis (DPA) plants. Ectopic expression of GhNAC8and GhNAC9show early leaf senescence compared to wild-types plant, revealed biological function for leaf senescence in Arabidopsis. The ten GhNAC genes displayed differential expression patterns and levels during natural and induced leaf senescence. Quantitative RT-PCR and promoter analyses suggest that these genes are induced by ABA, ethylene, drought, salinity, cold, heat and other hormonal treatments.
2. Expression analysis of NAC family gene (GhNAC18-77) for multiple abiotic stresses in cotton
A total of60full-length GhNAC genes were isolated from upland cotton, and they were phylogeneticly clustered into seven distinct subfamilies. We also constructed the phylogenetic tree for GhNAC genes and their motif, which show similarities among the subfamilies. The60GhNAC genes were located in the chromosome of D sub-genome and were found to be distributed on13chromosomes. Majority of the NACs showed specific temporal and spatial expression patterns for tissue-specific studies based on qRT-PCR analyses. Furthermore, the roles of GhNAC genes were monitored by qRT-PCR for leaf senescence, ethylene, ABA, GA3, drought and salinity.
3. GhNAC7, a novel GhNAC family transcription factor gene functions in leaf senescence
GhNAC7, a gene encoding a NAC family transcription factor was studied for functional analysis. Expression of this gene is closely associated with the leaf senescence process of cotton as revealed by tissue-specific and natural leaf senescence studies through qRT-PCR and immunoblot assays. The GhNAC7expression also induced by ethylene, ABA, drought, salinity, cold and high temperature treatments in cotton. In two model plant species, Arabidopsis and tobacco overexpression of GhNAC7showed visible symptoms for leaf senescence. Protein expression in BL21E. Coli and immunoblot assay in senescent leaf confirmed that molecular weight of GhNAC7protein was approximately29kDa. Our data strongly suggest that GhNAC7play an important role in cotton leaf senescence.
引文
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