陆地棉核雄性不育基因的分子标记定位及相关基因的克隆和功能分析
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摘要
棉花是一种重要的经济作物,在人们的生活中占有重要地位。棉花的杂种优势十分明显,可显著提高棉花的产量、品质、抗病和抗逆性。在生产上,棉花杂种优势利用主要是通过人工去雄杂交制种和利用棉花核雄性不育系杂交制种两种方法。利用核雄性不育,可以减少去雄用工,降低制种成本,所利用杂种一代集抗病、优质、高产等优点于一体,因而对棉花核雄性不育系的研究有着非常大的意义,在生产上有广泛的发展前景。因此,寻找到与核雄性不育(GMS)基因紧密连锁的分子标记将极大地推动GMS在棉花杂种优势上的应用。同时核雄性不育表达相关基因的克隆也将有助于了解雄性不育的作用机制,这无论对基础研究还是应用研究都具有重要意义。
本研究以陆地棉单隐性核雄性不育系阆A和重叠隐性核雄性不育中抗A为材料,分别构建了F_2和BC_1群体,通过分子标记技术对三个GMS基因进行了分子定位。以阆A的不育株(ms_(15)ms_(15))和可育株(Ms_(15)ms_(15))连续多代姐妹交培育而成的近等基因系的花药为材料,围绕小孢子减数分裂时期重点取材,利用差异显示技术克隆了一些可能与雄性不育表达的相关基因,并通过qRT-PCR验证了4个阳性差异片段。利用5′RACE和TAIL等方法克隆了一个差异片段的全长,命名为棉花过氧化酶基因Ghpod,该基因在花药中强优势表达,在营养器官中不表达。对另外一个差异基因Phi-1进行了转基因功能分析。目前取得如下研究进展:
1、棉花核雄性不育基因的分子标记定位
本研究利用SSR、SRAP等分子标记技术,对控制棉花核雄性不育系阆A和中抗A的三个核雄性不育基因ms_(15)、ms_5和ms_6进行了分子标记定位。
构建了(阆A×海7124)F_2群体,该群体大小为145株,育性分离比例符合3:1(x_c~2=2.21,0.10＜P＜0.25)。运用Joinmap将ms_(15)基因定位于棉花的第12号染色体中间部位。整个连锁群长度114.5 cM,标记间的平均距离为3.3 cM;ms_(15)被定位在NAU1278_270和NAU2176_240两个标记之间,遗传距离分别为1.9 cM和0.8 cM。共有13个SSR标记和2个SRAP标记与ms_(15)距离小于10.0 cM。
由于控制中抗A育性的是重叠隐性基因ms_5ms_6,直接定位ms_5ms_6有很大的难度。因此我们将(抗A×海7124)BC_1分离出的可育株再与中抗A不育株杂交,将符合3:1的BC_2家系对应的BC_1单株(基因型应该为Ms_5ms_5Ms_6ms_6)与BC_1中分离出的不育株重新构建成一个BC_1群体,用于定位ms_5╱ms_6基因,经卡方测验,这个新(抗A×海7124)BC_1群体也符合1:1(x_c~2=2.31,0.10＜P＜0.25)分离比例。运用Joinmap将ms_5定位在棉花第12染色体上,ms_6定位在第26染色体上。位于ms_5两侧标记为NAU3561,NAU2176和NAU2096,遗传距离分别为1.4 cM,1.8 cM和1.8 cM。位于ms_6两侧标记为BNL1227b和NAU460,遗传距离分别为1.5 cM和1.8 cM。
由于ms_5和ms_(15)被定位到棉花同一条染色体上,因此使用Joinmap整合功能构建了ms_5和ms_(15)的整合图谱,长度为117.0cM,平均标记间距离为3.34 cM;ms_(15)和ms_5在整合图谱上的距离为6.0cM,推测ms_5和ms_(15)可能不是等位基因。
本研究获得的与3个不育基因连锁的分子标记(多为SSR标记),可以应用于分子标记辅助选择育种,尤其是对重叠隐性核不育基因的分子定位,能够在早世代进行分子标记辅助选择,将大大提高育种效率。同时,对此3个核雄性不育基因的分子定位为图位克隆这3个不育基因奠定了基础。
2、DDRT-PCR技术克隆陆地棉阆A核雄性不育表达的相关基因
以陆地棉单隐性核雄性不育系阆A的不育株(ms_(15)ms_(15))与可育株(Ms_(15)ms_(15))的花药为材料,围绕小孢子减数分裂时期重点取材,利用差异显示技术(DDRT-PCR)克隆陆地棉阆A核雄性不育表达的相关基因,通过DDRT分析,获得了75个差异片段,对分子量大于150bp的65个片段进行测序及BLAST功能预测。通过对9个差异片段实时荧光定量PCR(qRT-PCR)分析,仅有4个基因符合DDRT结果。这4个阳性基因的BLAST功能预测结果为:
(1)2个与RNA结合蛋白(RNA-binding protein)基因相关:C7与富含氨基乙酸RNA结合蛋白(Glycine-rich RNA-binding protein)高度同源(97%);C17与含KH保守域RNA结合蛋白(KH-domain-containing protein)高度同源(93%)。RNA结合蛋白参与了RNA代谢的所有过程,包括细胞核加工、核-细胞质运输、稳定以及转录起始,RNA结合蛋白在基因调控过程中扮演了重要角色。在多细胞生物中,含KH保守域的蛋白往往参与转录、mRNA的稳定、翻译水平的沉默和mRNA的定位。含KH保守域的蛋白的突变往往会导致发育缺陷。(2)C103与钙结合蛋白(Calcium-binding protein)相关,可能与花粉管发育及细胞降解死亡有关。(3)C18与磷酸诱导蛋白基因Phi-1高度同源(99%)。
通过差异显示我们获得了一些可能与育性相关的片段,这些基因的功能需要进一步研究。
3、棉花过氧化酶基因Ghpod的克隆与特征分析
利用5′RACE和TAIL-PCR的方法,我们克隆了一个过氧化物酶基因,将其命名为Ghpod。该基因cDNA全长为1355bp,ORF为993bp,编码一个由330个氨基酸组成的蛋白;基因组全长为1588 bp,包含3个内含子和4个外显子。该基因翻译的蛋白包含保守的活性位点(66-77:AASLLRLHFHDC),一个近端的血红素配合基(194-204:DMIALSGAHTL),8个保守的半胱氨酸残基(C44,C77,C82,C124,C130,C209,C236,C326),组成4个2硫键(C44-C124,C77-C82,C130-C326,C209-236)(在http://www.expasy.ch/prosite中预测)。这些特征存在于绝大多数的ClassⅢ过氧化酶中。BLAST分析表明,该基因同血红素过氧化酶以及拟南芥过氧化酶具有很高的相似性。Ghpod氨基酸序列信号肽预测表明Ghpod具有一个22个氨基酸序列的信号肽(MAKCSVLLFLCLVMVSFNVANT),剪切位点介于Thr22和Met23之间。Ghpod与棉花中已报道的20个过氧化酶的多重比较显示Ghpod与它们的氨基酸序列相似性很低,介于28%和49%之间。RT-PCR和Northern表明该基因为花器官特异表达基因,在花药中强优势表达,在营养器官中不表达。Ghpod与花器官、花药发育的关系需要进一步的功能验证。Ghpod的特异表达模式表明该基因的启动子在花器官特异表达的调控机制研究中非常有用。Ghpod基因的克隆和表达特征的分析为我们研究生殖器官特异表达的植物过氧化酶提供了基础。
4、棉花Phi-1基因的克隆及功能分析
DDRT片段C18同棉花的Phosphate induced protein 1(Phi-1)基因同源性达99%,因而利用电子克隆的方法,获得了该基因的全长ORF。qRT-PCR分析结果显示Phi-1基因在花药发育的花粉母细胞减数分裂期至四分体时期,可育花药中的表达量高于不育花药中的表达量,尤其在减数分裂期差异显著,暗示Phi-1可能与棉花小孢子的败育有一定的相关性。Sano推测Phi-1基因是磷酸诱导型细胞周期特异性蛋白,在向因缺磷酸而导致的细胞周期停止的细胞中加入磷酸时,细胞周期重新启动,而Phi-1基因的表达也迅速启动。为了进一步研究Phi-1基因的功能,我们进行了转拟南芥功能验证。棉花Phi-1基因与拟南芥Phi-1基因的同源性比棉花Phi-1与Cell cycles-relatedgene(EXO)的同源性低,因而我们仅用构建的正义载体转拟南芥。T1、T2代过量表达Phi-1基因的植株育性正常,因而Phi-1基因的过量表达不能影响育性。过量表达Phi-1基因的植株的indole-3-pyruvate decarboxylase gene(ipdc)表达量明显低于野生型WT中的表达量,ipdc作为IAA合成的关键酶,我们推测ipdc表达量的降低可能导致IAA含量的减少,从而生长素IAA参与的植物生长的一系列过程可能受到影响。激素测定结果表明IAA含量明显降低,其它激素含量也有所变化。实验中我们观测到转基因的拟南芥主根长度比对照WT短、花期比对照WT晚,花序高度比对照WT低,可能原因是Phi-1基因的过量表达破坏了植物自身的细胞周期及影响了植物激素的表达,进而调节植物的细胞周期和发育进程。
Cotton is an important cash crop in the world and plays an important role in our life. The heterosis in cotton is very significant,especially in increasing yield,fiber quality and disease and stress resistance.The heterosis in practice production was mainly used by hand emasculation and male sterility.To use hand emasculation and pollination,utilization of male sterility is more effective and economical in producing commercial hybrid seeds,and can integrate the excellent traits of disease resistance,high quality and high yield in the hybrid F_1.Therefore,utilization of male sterility in heterosis has a good developmental prospect.An enhanced understanding of the genetics of GMS at the molecular level and the identification of linked molecular markers will greatly facilitate heterosis applications in cotton.The cloning of genes associated with the male sterility will also help to understand the mechanism of male sterility,which is greatly important to both basic and applied researches in cotton.
In this research,two populations,a F_2 and a new BC_1,were developed to map the ms_(15) and ms_5ms_6 male sterile genes.DDRT-PCR was performed between fertile and sterile pollens of Lang,A to look for male sterile related genes.To evaluate the accuraty of DDRT, 9 DDRT fragments were selected to perform Real-time quantitative PCR(qRT-PCR) and only 4 were positive fragments.A peroxidase gene,Ghpod,was cloned using 5'-RACE and TAIL-PCR,and was expressed in pollens and absent in vegetative tissues by RT-PCR and Northern analysis.Functional analysis of Phi-1 gene was performed by transgenic Arabidopsis.The results are following:
Molecular Mapping of male sterile genes ms_(15),ms_5 and ms_6:Three GMS genes,ms_(15)、ms_5 and ms_6,were mapped using SSR and SRAP in this research.
GMS line LangA was controlled by single recessive male sterile gene ms_(15) and ZhongkangA was controlled by duplicate gene,ms_5 and ms_6.The ms_(15) gene was mapped on chr.12,and was flanked by two SSR markers,NAU2176 and NAU1278,with a genetic distance of 0.8 cM and 1.9 cM,respectively.The ms_5 and ms_6 genes were mapped to one pair of homoeologous chromosomes,ms_5 on chr.12 flanked by three SSR markers, NAU3561,NAU2176 and NAU2096,with genetic distances of 1.4 cM,1.7 cM and 1.7 cM, respectively,and ms_6 on chr.26 flanked by two SSR markers,BNL1227 and NAU460,with a genetic distance of 1.3 cM and 1.7 cM,respectively.These markers were tightly linked with the ms_(15),ms_5 and ms_6 genes.Therefore they can be used in the marker-assisted selection of male-sterile plants among segregating populations in a breeding program,and provides the foundation for gene isolation by map-based cloning for these three genes.
Molecular cloning of male sterility related genes in upland cotton:Uing mRNA differential display,we obtained 65 cDNA fragments(＞150bp) differentially expressed in fertile and sterile pollen pools of isogenic lines ms_(15).To evaluate the accuraty of DDRT,9 DDRT fragments were selected to perform Real-time quantitative RT-PCR(qRT-PCR). Only 4 were basically associated with specificity found by DDRT.Sequencing and BLAST analysis showed that,among them there were 2 ESTs were homologous to RNA-binding protein:C7 was highly similar to Glycine-rich RNA-binding protein;C17 was highly similar to KH domain-containing protein.In metazoa,proteins containing KH domains have been implicated in transcription,mRNA stability,translational silencing and mRAN localization.Mutations of KH domain proteins usually result in developmental defects.We found two types of RNA-binding proteins in this study.The relation between them and the male sterility in cotton needed to be further studied.C103 was similar to Calcium-binding protein,which might be involved pollen tube development and apotosis in pollens.1 EST (C18) was highly similar to Phi-1 which was specific to phosphate-induced cell cycle activity,and played a role in causing phosphate-starved cells to re-enter the cell cycle.
We got some fragments which might be related to male sterility through DDRT-PCR technology,however the function of these fragments needed to be further investigated.
Moleculor cloning and charictorization of a flower-organ specific classⅢperoxidase gene in G.hirsutum:A cDNA clone encoding a 330 amina acid,designed Ghpod was isolated from the anthers of male sterility line LangA by using 5' RACE and TAIL-PCR technologies.Bioinform analysis indicated that Ghpod was high similarity to classⅢperoxidase.Ghpod has a signal peptide in the N terminal and a conserved active site residues,a proximal heme-ligand signature,eight conserved cysteine residues which yield the four disulfide bridges common in most classⅢperoxidase.Both the RT-PCR and Northern blots results showed that Ghpod was a flower-specific gene and predominantly expressed in pollen.The relationship between Ghpod and pollen development need further investigate.The Ghpod promoter will be useful for future studies of regulatory mechanisms controlling floral organ-specific expression.Specifically,it should be a useful tool for characterizing the spatial distribution patterns of transgenic pollen,to determine pollination mechanisms and to monitor gene flow in field conditions.
Over-expression of Phi-1 in transgenic Arabidopsis:The fragment of C18 got from DDRT has the 99%identity with cotton Phi-1 gene,so we obtained the full-length cDNA of this fragment using in silico cloning technique,qRT-PCR was performed to examine the Phi-1 expression pattern,qRT-PCR results demonstrated that Phi-1 was higher significantly expressed in fertile pollens development stages from meiosis to tetraspore stage than that of sterile pollens,especially in meiosis stages.It was supposed to be related to male sterility.It was reported that Phi-1 was specific to phosphate-induced cell cycle activity,and played a role in causing phosphate-starved ceils to re-enter the cell cycle.Based on the result that the sequence similarity between cotton Phi-1 and Arabidopsis Phi-1 was lower(62%) than its sequence similarity with Arabidopsis cell-cycle related protein(EXO)(70%),antisense might be not available to silence the gene in Arabidopsis.Therefore,the sense expression voctor of PBI121- Phi-1 were constructed to transfor Arabidopsis for further functional analysis of Phi-1.We got 2 T1 transgenic plants through Kan~+ selection and PCR identification.The T1 and T2 transgenic plants had no obvious change of pollen fertility. However,the expression of ipdc gene which was very important for IAA synthesis was decreased greatly in the over-expression transgenic Arabidopsis than in the WT Arabidosis. The comparative analysis of endogenous hormone contents in WT and transgenic Arabidopsis showed that IAA content was reduced obviously in transgenic Arabidopsis. The flowering time of T2 transgenic Arabidopsis was delayed about 6 days than that of WT. And the inflorescence/stem height of T2 transgenic Arabidopsis were reduced about 2.2 cM than that of WT.It was supposed that over-expression of Phi-1 in Arabidopsis might disturb the internal cell-cycle of plants and homeostasis of endogenous hormones,and therefore affected the growth and development of plants.
本研究以陆地棉单隐性核雄性不育系阆A和重叠隐性核雄性不育中抗A为材料,分别构建了F_2和BC_1群体,通过分子标记技术对三个GMS基因进行了分子定位。以阆A的不育株(ms_(15)ms_(15))和可育株(Ms_(15)ms_(15))连续多代姐妹交培育而成的近等基因系的花药为材料,围绕小孢子减数分裂时期重点取材,利用差异显示技术克隆了一些可能与雄性不育表达的相关基因,并通过qRT-PCR验证了4个阳性差异片段。利用5′RACE和TAIL等方法克隆了一个差异片段的全长,命名为棉花过氧化酶基因Ghpod,该基因在花药中强优势表达,在营养器官中不表达。对另外一个差异基因Phi-1进行了转基因功能分析。目前取得如下研究进展:
1、棉花核雄性不育基因的分子标记定位
本研究利用SSR、SRAP等分子标记技术,对控制棉花核雄性不育系阆A和中抗A的三个核雄性不育基因ms_(15)、ms_5和ms_6进行了分子标记定位。
构建了(阆A×海7124)F_2群体,该群体大小为145株,育性分离比例符合3:1(x_c~2=2.21,0.10＜P＜0.25)。运用Joinmap将ms_(15)基因定位于棉花的第12号染色体中间部位。整个连锁群长度114.5 cM,标记间的平均距离为3.3 cM;ms_(15)被定位在NAU1278_270和NAU2176_240两个标记之间,遗传距离分别为1.9 cM和0.8 cM。共有13个SSR标记和2个SRAP标记与ms_(15)距离小于10.0 cM。
由于控制中抗A育性的是重叠隐性基因ms_5ms_6,直接定位ms_5ms_6有很大的难度。因此我们将(抗A×海7124)BC_1分离出的可育株再与中抗A不育株杂交,将符合3:1的BC_2家系对应的BC_1单株(基因型应该为Ms_5ms_5Ms_6ms_6)与BC_1中分离出的不育株重新构建成一个BC_1群体,用于定位ms_5╱ms_6基因,经卡方测验,这个新(抗A×海7124)BC_1群体也符合1:1(x_c~2=2.31,0.10＜P＜0.25)分离比例。运用Joinmap将ms_5定位在棉花第12染色体上,ms_6定位在第26染色体上。位于ms_5两侧标记为NAU3561,NAU2176和NAU2096,遗传距离分别为1.4 cM,1.8 cM和1.8 cM。位于ms_6两侧标记为BNL1227b和NAU460,遗传距离分别为1.5 cM和1.8 cM。
由于ms_5和ms_(15)被定位到棉花同一条染色体上,因此使用Joinmap整合功能构建了ms_5和ms_(15)的整合图谱,长度为117.0cM,平均标记间距离为3.34 cM;ms_(15)和ms_5在整合图谱上的距离为6.0cM,推测ms_5和ms_(15)可能不是等位基因。
本研究获得的与3个不育基因连锁的分子标记(多为SSR标记),可以应用于分子标记辅助选择育种,尤其是对重叠隐性核不育基因的分子定位,能够在早世代进行分子标记辅助选择,将大大提高育种效率。同时,对此3个核雄性不育基因的分子定位为图位克隆这3个不育基因奠定了基础。
2、DDRT-PCR技术克隆陆地棉阆A核雄性不育表达的相关基因
以陆地棉单隐性核雄性不育系阆A的不育株(ms_(15)ms_(15))与可育株(Ms_(15)ms_(15))的花药为材料,围绕小孢子减数分裂时期重点取材,利用差异显示技术(DDRT-PCR)克隆陆地棉阆A核雄性不育表达的相关基因,通过DDRT分析,获得了75个差异片段,对分子量大于150bp的65个片段进行测序及BLAST功能预测。通过对9个差异片段实时荧光定量PCR(qRT-PCR)分析,仅有4个基因符合DDRT结果。这4个阳性基因的BLAST功能预测结果为:
(1)2个与RNA结合蛋白(RNA-binding protein)基因相关:C7与富含氨基乙酸RNA结合蛋白(Glycine-rich RNA-binding protein)高度同源(97%);C17与含KH保守域RNA结合蛋白(KH-domain-containing protein)高度同源(93%)。RNA结合蛋白参与了RNA代谢的所有过程,包括细胞核加工、核-细胞质运输、稳定以及转录起始,RNA结合蛋白在基因调控过程中扮演了重要角色。在多细胞生物中,含KH保守域的蛋白往往参与转录、mRNA的稳定、翻译水平的沉默和mRNA的定位。含KH保守域的蛋白的突变往往会导致发育缺陷。(2)C103与钙结合蛋白(Calcium-binding protein)相关,可能与花粉管发育及细胞降解死亡有关。(3)C18与磷酸诱导蛋白基因Phi-1高度同源(99%)。
通过差异显示我们获得了一些可能与育性相关的片段,这些基因的功能需要进一步研究。
3、棉花过氧化酶基因Ghpod的克隆与特征分析
利用5′RACE和TAIL-PCR的方法,我们克隆了一个过氧化物酶基因,将其命名为Ghpod。该基因cDNA全长为1355bp,ORF为993bp,编码一个由330个氨基酸组成的蛋白;基因组全长为1588 bp,包含3个内含子和4个外显子。该基因翻译的蛋白包含保守的活性位点(66-77:AASLLRLHFHDC),一个近端的血红素配合基(194-204:DMIALSGAHTL),8个保守的半胱氨酸残基(C44,C77,C82,C124,C130,C209,C236,C326),组成4个2硫键(C44-C124,C77-C82,C130-C326,C209-236)(在http://www.expasy.ch/prosite中预测)。这些特征存在于绝大多数的ClassⅢ过氧化酶中。BLAST分析表明,该基因同血红素过氧化酶以及拟南芥过氧化酶具有很高的相似性。Ghpod氨基酸序列信号肽预测表明Ghpod具有一个22个氨基酸序列的信号肽(MAKCSVLLFLCLVMVSFNVANT),剪切位点介于Thr22和Met23之间。Ghpod与棉花中已报道的20个过氧化酶的多重比较显示Ghpod与它们的氨基酸序列相似性很低,介于28%和49%之间。RT-PCR和Northern表明该基因为花器官特异表达基因,在花药中强优势表达,在营养器官中不表达。Ghpod与花器官、花药发育的关系需要进一步的功能验证。Ghpod的特异表达模式表明该基因的启动子在花器官特异表达的调控机制研究中非常有用。Ghpod基因的克隆和表达特征的分析为我们研究生殖器官特异表达的植物过氧化酶提供了基础。
4、棉花Phi-1基因的克隆及功能分析
DDRT片段C18同棉花的Phosphate induced protein 1(Phi-1)基因同源性达99%,因而利用电子克隆的方法,获得了该基因的全长ORF。qRT-PCR分析结果显示Phi-1基因在花药发育的花粉母细胞减数分裂期至四分体时期,可育花药中的表达量高于不育花药中的表达量,尤其在减数分裂期差异显著,暗示Phi-1可能与棉花小孢子的败育有一定的相关性。Sano推测Phi-1基因是磷酸诱导型细胞周期特异性蛋白,在向因缺磷酸而导致的细胞周期停止的细胞中加入磷酸时,细胞周期重新启动,而Phi-1基因的表达也迅速启动。为了进一步研究Phi-1基因的功能,我们进行了转拟南芥功能验证。棉花Phi-1基因与拟南芥Phi-1基因的同源性比棉花Phi-1与Cell cycles-relatedgene(EXO)的同源性低,因而我们仅用构建的正义载体转拟南芥。T1、T2代过量表达Phi-1基因的植株育性正常,因而Phi-1基因的过量表达不能影响育性。过量表达Phi-1基因的植株的indole-3-pyruvate decarboxylase gene(ipdc)表达量明显低于野生型WT中的表达量,ipdc作为IAA合成的关键酶,我们推测ipdc表达量的降低可能导致IAA含量的减少,从而生长素IAA参与的植物生长的一系列过程可能受到影响。激素测定结果表明IAA含量明显降低,其它激素含量也有所变化。实验中我们观测到转基因的拟南芥主根长度比对照WT短、花期比对照WT晚,花序高度比对照WT低,可能原因是Phi-1基因的过量表达破坏了植物自身的细胞周期及影响了植物激素的表达,进而调节植物的细胞周期和发育进程。
Cotton is an important cash crop in the world and plays an important role in our life. The heterosis in cotton is very significant,especially in increasing yield,fiber quality and disease and stress resistance.The heterosis in practice production was mainly used by hand emasculation and male sterility.To use hand emasculation and pollination,utilization of male sterility is more effective and economical in producing commercial hybrid seeds,and can integrate the excellent traits of disease resistance,high quality and high yield in the hybrid F_1.Therefore,utilization of male sterility in heterosis has a good developmental prospect.An enhanced understanding of the genetics of GMS at the molecular level and the identification of linked molecular markers will greatly facilitate heterosis applications in cotton.The cloning of genes associated with the male sterility will also help to understand the mechanism of male sterility,which is greatly important to both basic and applied researches in cotton.
In this research,two populations,a F_2 and a new BC_1,were developed to map the ms_(15) and ms_5ms_6 male sterile genes.DDRT-PCR was performed between fertile and sterile pollens of Lang,A to look for male sterile related genes.To evaluate the accuraty of DDRT, 9 DDRT fragments were selected to perform Real-time quantitative PCR(qRT-PCR) and only 4 were positive fragments.A peroxidase gene,Ghpod,was cloned using 5'-RACE and TAIL-PCR,and was expressed in pollens and absent in vegetative tissues by RT-PCR and Northern analysis.Functional analysis of Phi-1 gene was performed by transgenic Arabidopsis.The results are following:
Molecular Mapping of male sterile genes ms_(15),ms_5 and ms_6:Three GMS genes,ms_(15)、ms_5 and ms_6,were mapped using SSR and SRAP in this research.
GMS line LangA was controlled by single recessive male sterile gene ms_(15) and ZhongkangA was controlled by duplicate gene,ms_5 and ms_6.The ms_(15) gene was mapped on chr.12,and was flanked by two SSR markers,NAU2176 and NAU1278,with a genetic distance of 0.8 cM and 1.9 cM,respectively.The ms_5 and ms_6 genes were mapped to one pair of homoeologous chromosomes,ms_5 on chr.12 flanked by three SSR markers, NAU3561,NAU2176 and NAU2096,with genetic distances of 1.4 cM,1.7 cM and 1.7 cM, respectively,and ms_6 on chr.26 flanked by two SSR markers,BNL1227 and NAU460,with a genetic distance of 1.3 cM and 1.7 cM,respectively.These markers were tightly linked with the ms_(15),ms_5 and ms_6 genes.Therefore they can be used in the marker-assisted selection of male-sterile plants among segregating populations in a breeding program,and provides the foundation for gene isolation by map-based cloning for these three genes.
Molecular cloning of male sterility related genes in upland cotton:Uing mRNA differential display,we obtained 65 cDNA fragments(＞150bp) differentially expressed in fertile and sterile pollen pools of isogenic lines ms_(15).To evaluate the accuraty of DDRT,9 DDRT fragments were selected to perform Real-time quantitative RT-PCR(qRT-PCR). Only 4 were basically associated with specificity found by DDRT.Sequencing and BLAST analysis showed that,among them there were 2 ESTs were homologous to RNA-binding protein:C7 was highly similar to Glycine-rich RNA-binding protein;C17 was highly similar to KH domain-containing protein.In metazoa,proteins containing KH domains have been implicated in transcription,mRNA stability,translational silencing and mRAN localization.Mutations of KH domain proteins usually result in developmental defects.We found two types of RNA-binding proteins in this study.The relation between them and the male sterility in cotton needed to be further studied.C103 was similar to Calcium-binding protein,which might be involved pollen tube development and apotosis in pollens.1 EST (C18) was highly similar to Phi-1 which was specific to phosphate-induced cell cycle activity,and played a role in causing phosphate-starved cells to re-enter the cell cycle.
We got some fragments which might be related to male sterility through DDRT-PCR technology,however the function of these fragments needed to be further investigated.
Moleculor cloning and charictorization of a flower-organ specific classⅢperoxidase gene in G.hirsutum:A cDNA clone encoding a 330 amina acid,designed Ghpod was isolated from the anthers of male sterility line LangA by using 5' RACE and TAIL-PCR technologies.Bioinform analysis indicated that Ghpod was high similarity to classⅢperoxidase.Ghpod has a signal peptide in the N terminal and a conserved active site residues,a proximal heme-ligand signature,eight conserved cysteine residues which yield the four disulfide bridges common in most classⅢperoxidase.Both the RT-PCR and Northern blots results showed that Ghpod was a flower-specific gene and predominantly expressed in pollen.The relationship between Ghpod and pollen development need further investigate.The Ghpod promoter will be useful for future studies of regulatory mechanisms controlling floral organ-specific expression.Specifically,it should be a useful tool for characterizing the spatial distribution patterns of transgenic pollen,to determine pollination mechanisms and to monitor gene flow in field conditions.
Over-expression of Phi-1 in transgenic Arabidopsis:The fragment of C18 got from DDRT has the 99%identity with cotton Phi-1 gene,so we obtained the full-length cDNA of this fragment using in silico cloning technique,qRT-PCR was performed to examine the Phi-1 expression pattern,qRT-PCR results demonstrated that Phi-1 was higher significantly expressed in fertile pollens development stages from meiosis to tetraspore stage than that of sterile pollens,especially in meiosis stages.It was supposed to be related to male sterility.It was reported that Phi-1 was specific to phosphate-induced cell cycle activity,and played a role in causing phosphate-starved ceils to re-enter the cell cycle.Based on the result that the sequence similarity between cotton Phi-1 and Arabidopsis Phi-1 was lower(62%) than its sequence similarity with Arabidopsis cell-cycle related protein(EXO)(70%),antisense might be not available to silence the gene in Arabidopsis.Therefore,the sense expression voctor of PBI121- Phi-1 were constructed to transfor Arabidopsis for further functional analysis of Phi-1.We got 2 T1 transgenic plants through Kan~+ selection and PCR identification.The T1 and T2 transgenic plants had no obvious change of pollen fertility. However,the expression of ipdc gene which was very important for IAA synthesis was decreased greatly in the over-expression transgenic Arabidopsis than in the WT Arabidosis. The comparative analysis of endogenous hormone contents in WT and transgenic Arabidopsis showed that IAA content was reduced obviously in transgenic Arabidopsis. The flowering time of T2 transgenic Arabidopsis was delayed about 6 days than that of WT. And the inflorescence/stem height of T2 transgenic Arabidopsis were reduced about 2.2 cM than that of WT.It was supposed that over-expression of Phi-1 in Arabidopsis might disturb the internal cell-cycle of plants and homeostasis of endogenous hormones,and therefore affected the growth and development of plants.
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