玉米匍匐基因LAZY1的图位克隆及功能分析
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摘要
生长素在植物生长发育的各个方面均发挥重要作用,包括重力性生长和花器官发育。尽管目前已分别鉴定了多个重力性相关或花器官发育相关的基因,但植物体内尚未有明确报道同时参与这两个过程的基因。玉米经典的遗传学突变体之一lazy plantl (lal)发现几十年来,人们早已观察到其地上部重力性缺失的突变表型特征,然而LA1基因调控玉米地上部重力性生长的分子机理却一直未有研究。
本研究中,我们利用分子遗传分析、发育生物学手段及转录组分析对玉米LAl基因进行了功能剖析,发现其为水稻和拟南芥LAZY1基因的同源基因,在玉米中特异调控地上部重力性反应和花器官发育。主要研究结果如下：
1)自MuDR突变体库中筛选获得匍匐生长突变体prostrate steml (ps1),通过图位克隆策略获得候选目的基因。等位测验发现psl突变体是玉米la1的等位突变,表现为地上部重力性缺失而造成茎杆匍匐生长。与水稻或拟南芥同源基因不同,玉米LA1基因除在茎节中表达外还在生殖器官中表达,且参与调控花器官发育。
2)LA1转录对外源生长素和光信号敏感。施加生长素后可诱导LA1的转录水平先下调后上调。黑暗环境诱导LA1基因表达而光照抑制该基因表达。
3)LA1蛋白定位于细胞核和细胞膜上；通过酵母双杂交和双分子荧光互补(BiFC)实验鉴定了LA1蛋白可与一个蛋白激酶C (PKC)在膜上互作,可能作用于生长素运输的调控过程；还可与一个AUX/IAA蛋白(IAA17)在细胞核内互作,可能作用于生长素的信号转导途径。
4)转录组测序分析显示部分生长素运输载体或相关基因,如ZmPIN1c,在la1突变体中表达量明显降低；而另有部分运输载体及相关基因,如BR2(ZmABCB1),则明显上调表达。此结果与LA1对生长素极性运输的双向调控作用是一致的：负向调控向基极性运输而正向调控侧向极性运输。另外,la1突变体中大量的生长素响应基因和光信号相关基因发生不同程度的上调变化,预示LA1在两个过程中的抑制调节作用。
综上,本研究证明LA1参与玉米生长素极性运输、生长素信号转导及光信号响应等一系列基础的生命过程,介导了重力性生长和花器官形态建成之间的互作。相关结果不仅增进了我们对生长素作用于玉米重力性生长、花结构发育和光形态建成间紧密关系的了解,同时也将为未来玉米抗倒伏及理想株型的选育提供理论依据。
The plant hormone auxin plays important role in multitude of physiological and developmental processes including shoot gravitropism and inflorescence development. Despite many genes have been characterized to be involved in either process, few regulators, if any, have been identified to mediate both gravitropism and inflorescence development in a given species. The classical maize mutant lazy plantl (laI) have been known for decades to have defected gravitropic response, yet the mechanism underlining maize gravitropism remain largely elusive.
In this research, we combined genetic analyses, developmental techniques, biochemical tools with transcriptome analyses to identify LAI, an ortholog of LAZY1in rice and in Arabidopsis, regulating both shoot gravitropism and inflorescence development in maize. The major novel discoveries of our study are:
1) We created a maize prostratel (psl) mutant from our MuDR mutant library, which is allelic to the classic lal mutant, displaying a prostrate phenotype with reduced shoot gravitropism. We map-based cloned the LAI gene. Unlike its orthologs in rice or Arabidopsis, maize LAI regulates maize inflorescence development, and consistently, LAI is highly expressed in the reproductive organs.
2) Transcription of LAI is responsive to both auxin and light. Exogenous auxin treatment led to a repression-then-induction pattern for LAI expression, while dark dramatically induced the transcript accumulation of LAI and light inhibited it.
3) LA1protein localizes to the plasma membrane as well as in the nucleus. LA1physically interacts with a putative auxin transport regulator (Protein Kinase C, PKC) in the plasma membrane and a putative auxin signaling protein (AUX/IAA protein IAA17) in the nucleus, linking LA1with auxin transport and auxin signaling.
4) RNA-SEQ analyses of lal mutant node showed auxin transport related genes, such as ZmPIN1c, were greatly downregulated while maize BR2(ZmABCBl) was significantly upregulated, consistant with the bilateral function of LA1of suppressing the basipetal polar auxin transport (PAT) while promoting the lateral PAT in maize. Further, most auxin responsive elements and light signaling genes were differentially up-regulated in the lal nodes, implying that LA1may act as a negative regulator for both light signaling and auxin signaling.
Taken together, our findings suggest that LA1may mediate the crosstalk between shoot gravitropism and inflorescence development by regulating auxin transport, auxin signaling and probably light response in maize. As such, the results not only advance our understanding of the function of LA1in response to gravity, but also give new insights into the complicated crosstalk between auxin, light and inflorescent development.
本研究中,我们利用分子遗传分析、发育生物学手段及转录组分析对玉米LAl基因进行了功能剖析,发现其为水稻和拟南芥LAZY1基因的同源基因,在玉米中特异调控地上部重力性反应和花器官发育。主要研究结果如下：
1)自MuDR突变体库中筛选获得匍匐生长突变体prostrate steml (ps1),通过图位克隆策略获得候选目的基因。等位测验发现psl突变体是玉米la1的等位突变,表现为地上部重力性缺失而造成茎杆匍匐生长。与水稻或拟南芥同源基因不同,玉米LA1基因除在茎节中表达外还在生殖器官中表达,且参与调控花器官发育。
2)LA1转录对外源生长素和光信号敏感。施加生长素后可诱导LA1的转录水平先下调后上调。黑暗环境诱导LA1基因表达而光照抑制该基因表达。
3)LA1蛋白定位于细胞核和细胞膜上；通过酵母双杂交和双分子荧光互补(BiFC)实验鉴定了LA1蛋白可与一个蛋白激酶C (PKC)在膜上互作,可能作用于生长素运输的调控过程；还可与一个AUX/IAA蛋白(IAA17)在细胞核内互作,可能作用于生长素的信号转导途径。
4)转录组测序分析显示部分生长素运输载体或相关基因,如ZmPIN1c,在la1突变体中表达量明显降低；而另有部分运输载体及相关基因,如BR2(ZmABCB1),则明显上调表达。此结果与LA1对生长素极性运输的双向调控作用是一致的：负向调控向基极性运输而正向调控侧向极性运输。另外,la1突变体中大量的生长素响应基因和光信号相关基因发生不同程度的上调变化,预示LA1在两个过程中的抑制调节作用。
综上,本研究证明LA1参与玉米生长素极性运输、生长素信号转导及光信号响应等一系列基础的生命过程,介导了重力性生长和花器官形态建成之间的互作。相关结果不仅增进了我们对生长素作用于玉米重力性生长、花结构发育和光形态建成间紧密关系的了解,同时也将为未来玉米抗倒伏及理想株型的选育提供理论依据。
The plant hormone auxin plays important role in multitude of physiological and developmental processes including shoot gravitropism and inflorescence development. Despite many genes have been characterized to be involved in either process, few regulators, if any, have been identified to mediate both gravitropism and inflorescence development in a given species. The classical maize mutant lazy plantl (laI) have been known for decades to have defected gravitropic response, yet the mechanism underlining maize gravitropism remain largely elusive.
In this research, we combined genetic analyses, developmental techniques, biochemical tools with transcriptome analyses to identify LAI, an ortholog of LAZY1in rice and in Arabidopsis, regulating both shoot gravitropism and inflorescence development in maize. The major novel discoveries of our study are:
1) We created a maize prostratel (psl) mutant from our MuDR mutant library, which is allelic to the classic lal mutant, displaying a prostrate phenotype with reduced shoot gravitropism. We map-based cloned the LAI gene. Unlike its orthologs in rice or Arabidopsis, maize LAI regulates maize inflorescence development, and consistently, LAI is highly expressed in the reproductive organs.
2) Transcription of LAI is responsive to both auxin and light. Exogenous auxin treatment led to a repression-then-induction pattern for LAI expression, while dark dramatically induced the transcript accumulation of LAI and light inhibited it.
3) LA1protein localizes to the plasma membrane as well as in the nucleus. LA1physically interacts with a putative auxin transport regulator (Protein Kinase C, PKC) in the plasma membrane and a putative auxin signaling protein (AUX/IAA protein IAA17) in the nucleus, linking LA1with auxin transport and auxin signaling.
4) RNA-SEQ analyses of lal mutant node showed auxin transport related genes, such as ZmPIN1c, were greatly downregulated while maize BR2(ZmABCBl) was significantly upregulated, consistant with the bilateral function of LA1of suppressing the basipetal polar auxin transport (PAT) while promoting the lateral PAT in maize. Further, most auxin responsive elements and light signaling genes were differentially up-regulated in the lal nodes, implying that LA1may act as a negative regulator for both light signaling and auxin signaling.
Taken together, our findings suggest that LA1may mediate the crosstalk between shoot gravitropism and inflorescence development by regulating auxin transport, auxin signaling and probably light response in maize. As such, the results not only advance our understanding of the function of LA1in response to gravity, but also give new insights into the complicated crosstalk between auxin, light and inflorescent development.
引文
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