水稻T-DNA插入突变体侧翼序列的分离与分析及控制杂合单株低育性基因Osfbox的功能研究
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摘要
创造基因功能缺失突变体是研究目的基因功能的最直接和有效的途径之一。农杆菌介导的T-DNA插入不仅破坏了插入位点的基因功能,而且因为7-DNA序列已知,使得被破坏的基因序列的分离相对简单和快捷。农杆菌介导的T-DNA转化还因其在水稻基因组中相对随机的分布规律,成熟的转化体系,稳定的后代遗传,简单的遗传背景,被广泛地用于水稻插入突变体库的构建。
T-DNA侧翼序列的分离对水稻T-DNA插入突变体库的应用至关重要。一方面,我们可以根据大量的T-DNA侧翼序列来分析T-DNA在水稻基因组中的分布规律,进而明晰农杆菌介导的T-DNA转化机制;另一方面,根据侧翼序列提供的基因信息,我们可以广泛地开展反向遗传学的研究。本研究中,我们采用PCR扩增对突变体的阳性情况进行了检测,利用TAIL-PCR的方法成功地分离T-DNA侧翼序列,分析了T-DNA在水稻基因组中的分布规律,揭示了T-DNA插入位点序列的DNA物理特性和碱基组成特征。根据已有的侧翼序列,我们还开展了反向遗传学的研究,得到了一个杂合低育性的突变体,在细胞学水平观察了突变体败育的原因,并对控制突变性状的候选目标基因Osfbox的功能进行了详细的分析,主要研究结果如下:
1.以T-DNA上的GAL4/VP16基因区段为标记,对9024个转基因植株DNA进行了PCR扩增阳性检测,扩增结果表明7862为PCR阳性,阳性率为87.1%。
2.利用TAIL-PCR的方法,本人成功的分离得到了5100条水稻T-DNA侧翼序列,其中2609条可以准确的定位在水稻基因组上,定位率为51.2%。联合课题组其它五位同学的侧翼序列,课题组共有侧翼序列30577条,定位率为51.5%。
3.无论是从T-DNA标签的绝对数量还是相对密度来看,T-DNA都偏爱于插入到水稻中长度较大的染色体上,且插入密度与染色体大小显著正相关(r=0.74,P＜0.01)。在同一染色体上,T-DNA的分布更倾向于插入到远离着丝粒的染色体末端,且与染色体区段上的全长cDNA数目显著相关(r=0.70,P＜0.01)。T-DNA在水稻基因组中极度地偏向于不插入到转座子相关序列中(SR=—34.0)。在水稻的基因区域中,我们发现T-DNA偏爱于插入到上游1kb区(SR=12.8)和下游500bp区(SR=10.3)而趋向于不插入到编码区(SR=—13.2).但在基因的编码区内,我们并没有发现T-DNA在外显子和内含子之间存在偏爱性(χ~2=2.81,P=0.09)。
4.T-DNA在不同“分子功能”的基因中同样也存在着分布的偏爱性(χ~2=93.5,P＜0.000001);偏向于插入到“Antioxidant”(SR=6.07)和“Catalytic”(SR=3.79)两类功能基因中,偏向于不插入到“Nutrient reservoir”,“Enzymeregulator”,“Transcription regulator”和“Ligand binding and carrier”(SR值介于—2.1到—3.8),而在其它的五类功能基因中基本上呈一种随机分布的状态。
5.我们分析了本室和来源于其它三家研究单位的水稻T-DNA插入位点前后各1kb(共2001bp)序列(ISNS)的弯曲度的平均值,并以随机提取的2000条长度在2001bp的序列作为对照。弯曲度图的结果显示:随机序列的弯曲度值表现为杂乱的曲线。而本室提取的ISNS的弯曲度图中我们可以看到在序列的—200bp到200bp这一序列区间内弯曲度值出现了一个明显的波型变化,在—200和200位置为波谷。最大值波峰分别出现在约—10和10的位置,且在0,即T-DNA插入位点陡然的下降出现一个相对低值.在其它研究单位提取的ISNS中我们同样观察到了同样的现象。
6.对不同ISNS的GC含量分析后我们认为GC含量的高低不是T-DNA插入位点的主要特征。但对ISNS和随机提取序列的GC skew和TA skew分析发现:两组ISNS的GC与ZA skews之间呈显著的负相关(r＞-0.92).GC与TA skew曲线在插入位点位置出现交叉,且两种skew的值在插入位点都约为0;在-800到0的这段序列里,GC skew值表现为正值,且在-200到-100位置达到最大值,与之对应的是TA skew值为负值,但也是在同样位置达到最小值;在0-800区域内我们也发现了对应的结果,GC skew值表现为负值,TA skew值表现为正值,且在200到100位置碱基的分布出现极度的不均衡情况。在对照序列的结果中并没有发现这些特征。
7.在我们的T-DNA插入群体中,同一插入位点的T-DNA串连重复普遍存在,正向重复,头对头反向重复和尾对尾反向重复的比例分别达到了19.9%,10.5%和25.4%,40.9%的单株中至少存在着一种以上的串连重复形式;载体骨架序列带入到水稻基因组中的情况同样突出,左右边界带入的载体骨架序列单株比例分别达到了49.6%和29.7%。
8.对一个T-DNA插入位点位于第六染色体上的f-box基因3'UTR内的突变体开展了反向遗传学的研究,发现T-DNA杂合单株表现为半不育,但T-DNA阴性和纯合单株均育性正常,类似于水稻籼粳亚种间杂种不育。PCR扩增结果显示突变表型与T-DNA插入位点满足严格的共分离关系。目标基因被命名为Osfbox。
9。Osfbox在TIGR中的编号为LOC_Os06g06050,基因全长4441bp。KOME提供了该基因的全长cDNA信息(AK065478):cDNA的总长度为3723bp,由4个外显子和3个内含子组成,编码一个包含720个氨基酸的蛋白,在第15—63氨基酸位置存在一个F-box domain,390—469氨基酸位置为Cysteine-containing LRR profile结构域。
10.细胞学观察结果发现雌雄配子发育的受阻共同导致了T-DNA杂合单株的低育性。我们在小孢子的四分体时期开始发现绒毡层的降解出现了滞后。大、均能通过减数分裂时期,但在随后的有丝分裂过程中受阻,最(?)小孢子母细终导致败育。
11.对10个典型的水稻籼粳品种的Osfbox基因的序列比对结果表明在籼粳品种之间的碱基变异共有18处,其中12处发生在基因转录起始位点的前648bp范围内。在基因的编码区,我们只发现了粳稻基因型存在2处缺失,导致两个位点粳稻的氨基酸序列分别产生了3个串连谷氨酸和1个甘氨酸的缺失。
12.全生育期表达谱芯片和RT-PCR结果显示Osfbox是一个组成型表达基因,在雌雄蕊中的表达量稍高。RT-PCR的结果还表明在三到八期幼穗中,四期的幼穗中Osfbox的表达量达到最高,尔后随着时间的推移逐步减弱。原位杂交的结果也显示直至小孢子母细胞形成四分体,Osfbox都有较强烈的表达,但液泡化花粉期后花药中基本检测不到基因的表达。
13.Osfbox基因编码的蛋白专一地定位在细胞核中。
Generating loss-of-function mutant is one of the most strait-forward and efficient way to study gene's bio-function.Agro-bacterium induced T-DNA integration not only disrupt the gene function of the insertion site,but also facilitate the gene cloning for the highly developed methods in isolating the sequences flanking the T-DNA. With a relatively random distribution pattern in rice genome,highly efficient transformation system,stable inheritance in progeny and simple genetic background, Agro-bacterium induced T-DNA transformation has been widely used in rice insertional mutant library establishment.
The isolation of the flanking sequence tags(FST) is of great importance in the application of rice T-DNA insertional mutant library.On one hand,based on the vast FST data,we can have a deep insight into the T-DNA distribution pattern in rice genome and then elucidate the mechanism of Agro-bacterium induced T-DNA transformation;on the other hand,we can carry out reverse genetics study of the corresponding genes according to,the tagged gene information.
In this study,I accounted the transformation positive ratio of our insertional population by PCR marker in T-DNA.By employing the TAIL-PCR method,30577 T-DNA FSTs were successfully isolated;T-DNA distribution pattern was analyzed, and several DNA physical property and base composition features of the insertion sites were revealed.With the tagged gene information,reverse genetics study was carried out here and a mutant which showed a semi-sterile phenotype in hemizygous lines was obtained.Cytological analysis of the mutant and functional analysis of the candidate gene Osfbox have been conducted.The main results are described as follows:
1.The transformation positive ratio of 9024 transformants was checked by using GAL4/VP16 fragment in T-DNA as a PCR marker,7862(87.1%) showed to be PCR positive.
2.Employing the TAIL-PCR method,I independently isolated 5100 FSTs,2609 (51.2%) out of which can be well mapped onto the dee genome.Combined with the FSTs isolated by colleagues,there are totally 30577 FSTs in our group, including 51.5%can be mapped.
3.T-DNA insertions were biased towards large chromosomes,not only in the absolute number of insertions but also in the relative density,and the insertion density is highly correlated with the chromosome size(r= 0.74,P＜0.01).Within chromosomes,the insertions occurred more densely in the distal ends,and less densely in the centromeric regions and the insertion number is significantly correlated with the cDNA number in corresponding region(r=0.70,P＜0.01); T-DNA insertions strongly disfavored transposable element(TE)-related sequences(SR=—34.0) and favored genic sequences(SR=15.55).In rice genes, strong bias toward the 5' upstream(SR=12.8)and Y downstream regions(SR=10.3) of the genes were detected respectively,but the coding sequence region is observed to be disfavored(SR=—13.2).In the coding region,no insertional bias between intron and exon region was observed(χ~2 = 2.81,P = 0.09).
4.According to the "molecular functions" of genes,T-DNA insertions bias among the various classes of functional genes were observed(χ~2=93.5,P＜0.000001): preferentially occurred in "Antioxidant"(SR =6.07)and "Catalytic"(SR=3.79), but preferentially not occurred in "Nutrient reservoir","Enzyme regulator", "Transcription regulator" and "Ligand binding and cartier"(SR between -2.1 to -3.8).In the other 5 gene classes,T-DNA showed a random distribution pattern.
5.In ISNS bendability analysis result,the random CK showed an irregular curve. But in our ISNS,elevated bendability was observed at positions from -200 to 200 bp and the bendability peak is symmetric in this region,with the highest points at both -10 and 10 bp from the insertion sites.There was an abrupt drop in the bendability value at the insertion sites.A similar result was also observed in ISNS from other 3 research groups.
6.The analysis of ISNS GC content indicated that GC content is not a necessary factor for T-DNA insertion site.But The GC skew and TA skew calculation exhibited several features in both ISNS from our lab and 3 other groups:The GC and TA skews appeared to be inversely correlated with r＞-0.92;The two curves crossed each other at the insertion sites(point 0) at which both GC and TA skews were equal to 0;From the insertion site to 800-bp upstream,the GC skew was positive and reached a plateau in approximately the region from -300 to -100 bp, whereas the TA skew was negative and formed a valley in a similar position in approximately the region from -300 to -100 bp.The reverse was the case from the insertion sites to 800 bp downstream,whereas no such features were observed in the 2000 random sequences.
7.In our T-DNA insertional population,T-DNA inserted in one loci as a "tandem repeats" was widely detected.The ratio of "direct repeat","5' end junction inverse repeat" and "3' end junction inverse repeat" can be up to 19.9%,10.5% and 25.4%respectively.40.9%are detected to have at least one tandem repeat. The introduction of vector backbone sequence into the rice genome are prominent as well,up to 49.6%and 29.7%of the transformants habor a backbone sequence in left and right border respectively.
8.Reverse genetics study was carried out in a mutant which harboring a T-DNA insertion in the 3'UTR of an f-box gene in chromosome 6.The T-DNA hemizygous lines are semi-sterile,but both the T-DNA negative and homozygous lines are full fertile.This is similar to the semi-sterility of indica-japonica subspecies hybrids.The mutant phenotype is checked to be co-segregated with the T-DNA insertion.The candidate gene is named as Osfoox.
9.The ID of Osfoox in TIGR is LOC_Os06g06050 with a length of 4441bp.The full-length eDNA information in KOME(AK065478) showed that:3723bp in length,containes 4 exons and 3 introns,encoding a 720AA protein.A potential F-box domain in 15-63 AA and Cysteine-eontaining LRR profile domain in 390-469AA were detected by Plantsp.
10.Cytological analysis showed that tapetum degeneration retardation happened in tetrad stage,and the semi-sterility of T-DNA hemizygous lines are a consequence of both the undeveloped microspore and megaspore,which can pass the meiosis stage but are blocked in the following mitosis stage.
11.By comparing the Osfbox gene sequence of 10 typical rice indica or japonica varieties,we found 18 base variations between the 2 subspecies,including 12 occurred in the 648bp upstream region from the transcription start site.In gene coding region,only 2 deletions were detected,resulting in a 3 tandem Glu and 1 Gly deletion respectively.
12.Microarray based expression profile and RT-PCR data showed that Osfbox is a constitutively expressed gene.A higher expression level was detected in anther and pistil.In young panicle development stage 3-8,Osfbox expression level reaches the peak in stage 4 and then decreased.In situ hybridization result also showed that Osfbox is strongly expressed until tetrad formatted,but after the vacuolated pollen stage,no expression was detected in pollens.
13.The Osfbox protein is specifically localized in cell nuclear.
T-DNA侧翼序列的分离对水稻T-DNA插入突变体库的应用至关重要。一方面,我们可以根据大量的T-DNA侧翼序列来分析T-DNA在水稻基因组中的分布规律,进而明晰农杆菌介导的T-DNA转化机制;另一方面,根据侧翼序列提供的基因信息,我们可以广泛地开展反向遗传学的研究。本研究中,我们采用PCR扩增对突变体的阳性情况进行了检测,利用TAIL-PCR的方法成功地分离T-DNA侧翼序列,分析了T-DNA在水稻基因组中的分布规律,揭示了T-DNA插入位点序列的DNA物理特性和碱基组成特征。根据已有的侧翼序列,我们还开展了反向遗传学的研究,得到了一个杂合低育性的突变体,在细胞学水平观察了突变体败育的原因,并对控制突变性状的候选目标基因Osfbox的功能进行了详细的分析,主要研究结果如下:
1.以T-DNA上的GAL4/VP16基因区段为标记,对9024个转基因植株DNA进行了PCR扩增阳性检测,扩增结果表明7862为PCR阳性,阳性率为87.1%。
2.利用TAIL-PCR的方法,本人成功的分离得到了5100条水稻T-DNA侧翼序列,其中2609条可以准确的定位在水稻基因组上,定位率为51.2%。联合课题组其它五位同学的侧翼序列,课题组共有侧翼序列30577条,定位率为51.5%。
3.无论是从T-DNA标签的绝对数量还是相对密度来看,T-DNA都偏爱于插入到水稻中长度较大的染色体上,且插入密度与染色体大小显著正相关(r=0.74,P＜0.01)。在同一染色体上,T-DNA的分布更倾向于插入到远离着丝粒的染色体末端,且与染色体区段上的全长cDNA数目显著相关(r=0.70,P＜0.01)。T-DNA在水稻基因组中极度地偏向于不插入到转座子相关序列中(SR=—34.0)。在水稻的基因区域中,我们发现T-DNA偏爱于插入到上游1kb区(SR=12.8)和下游500bp区(SR=10.3)而趋向于不插入到编码区(SR=—13.2).但在基因的编码区内,我们并没有发现T-DNA在外显子和内含子之间存在偏爱性(χ~2=2.81,P=0.09)。
4.T-DNA在不同“分子功能”的基因中同样也存在着分布的偏爱性(χ~2=93.5,P＜0.000001);偏向于插入到“Antioxidant”(SR=6.07)和“Catalytic”(SR=3.79)两类功能基因中,偏向于不插入到“Nutrient reservoir”,“Enzymeregulator”,“Transcription regulator”和“Ligand binding and carrier”(SR值介于—2.1到—3.8),而在其它的五类功能基因中基本上呈一种随机分布的状态。
5.我们分析了本室和来源于其它三家研究单位的水稻T-DNA插入位点前后各1kb(共2001bp)序列(ISNS)的弯曲度的平均值,并以随机提取的2000条长度在2001bp的序列作为对照。弯曲度图的结果显示:随机序列的弯曲度值表现为杂乱的曲线。而本室提取的ISNS的弯曲度图中我们可以看到在序列的—200bp到200bp这一序列区间内弯曲度值出现了一个明显的波型变化,在—200和200位置为波谷。最大值波峰分别出现在约—10和10的位置,且在0,即T-DNA插入位点陡然的下降出现一个相对低值.在其它研究单位提取的ISNS中我们同样观察到了同样的现象。
6.对不同ISNS的GC含量分析后我们认为GC含量的高低不是T-DNA插入位点的主要特征。但对ISNS和随机提取序列的GC skew和TA skew分析发现:两组ISNS的GC与ZA skews之间呈显著的负相关(r＞-0.92).GC与TA skew曲线在插入位点位置出现交叉,且两种skew的值在插入位点都约为0;在-800到0的这段序列里,GC skew值表现为正值,且在-200到-100位置达到最大值,与之对应的是TA skew值为负值,但也是在同样位置达到最小值;在0-800区域内我们也发现了对应的结果,GC skew值表现为负值,TA skew值表现为正值,且在200到100位置碱基的分布出现极度的不均衡情况。在对照序列的结果中并没有发现这些特征。
7.在我们的T-DNA插入群体中,同一插入位点的T-DNA串连重复普遍存在,正向重复,头对头反向重复和尾对尾反向重复的比例分别达到了19.9%,10.5%和25.4%,40.9%的单株中至少存在着一种以上的串连重复形式;载体骨架序列带入到水稻基因组中的情况同样突出,左右边界带入的载体骨架序列单株比例分别达到了49.6%和29.7%。
8.对一个T-DNA插入位点位于第六染色体上的f-box基因3'UTR内的突变体开展了反向遗传学的研究,发现T-DNA杂合单株表现为半不育,但T-DNA阴性和纯合单株均育性正常,类似于水稻籼粳亚种间杂种不育。PCR扩增结果显示突变表型与T-DNA插入位点满足严格的共分离关系。目标基因被命名为Osfbox。
9。Osfbox在TIGR中的编号为LOC_Os06g06050,基因全长4441bp。KOME提供了该基因的全长cDNA信息(AK065478):cDNA的总长度为3723bp,由4个外显子和3个内含子组成,编码一个包含720个氨基酸的蛋白,在第15—63氨基酸位置存在一个F-box domain,390—469氨基酸位置为Cysteine-containing LRR profile结构域。
10.细胞学观察结果发现雌雄配子发育的受阻共同导致了T-DNA杂合单株的低育性。我们在小孢子的四分体时期开始发现绒毡层的降解出现了滞后。大、均能通过减数分裂时期,但在随后的有丝分裂过程中受阻,最(?)小孢子母细终导致败育。
11.对10个典型的水稻籼粳品种的Osfbox基因的序列比对结果表明在籼粳品种之间的碱基变异共有18处,其中12处发生在基因转录起始位点的前648bp范围内。在基因的编码区,我们只发现了粳稻基因型存在2处缺失,导致两个位点粳稻的氨基酸序列分别产生了3个串连谷氨酸和1个甘氨酸的缺失。
12.全生育期表达谱芯片和RT-PCR结果显示Osfbox是一个组成型表达基因,在雌雄蕊中的表达量稍高。RT-PCR的结果还表明在三到八期幼穗中,四期的幼穗中Osfbox的表达量达到最高,尔后随着时间的推移逐步减弱。原位杂交的结果也显示直至小孢子母细胞形成四分体,Osfbox都有较强烈的表达,但液泡化花粉期后花药中基本检测不到基因的表达。
13.Osfbox基因编码的蛋白专一地定位在细胞核中。
Generating loss-of-function mutant is one of the most strait-forward and efficient way to study gene's bio-function.Agro-bacterium induced T-DNA integration not only disrupt the gene function of the insertion site,but also facilitate the gene cloning for the highly developed methods in isolating the sequences flanking the T-DNA. With a relatively random distribution pattern in rice genome,highly efficient transformation system,stable inheritance in progeny and simple genetic background, Agro-bacterium induced T-DNA transformation has been widely used in rice insertional mutant library establishment.
The isolation of the flanking sequence tags(FST) is of great importance in the application of rice T-DNA insertional mutant library.On one hand,based on the vast FST data,we can have a deep insight into the T-DNA distribution pattern in rice genome and then elucidate the mechanism of Agro-bacterium induced T-DNA transformation;on the other hand,we can carry out reverse genetics study of the corresponding genes according to,the tagged gene information.
In this study,I accounted the transformation positive ratio of our insertional population by PCR marker in T-DNA.By employing the TAIL-PCR method,30577 T-DNA FSTs were successfully isolated;T-DNA distribution pattern was analyzed, and several DNA physical property and base composition features of the insertion sites were revealed.With the tagged gene information,reverse genetics study was carried out here and a mutant which showed a semi-sterile phenotype in hemizygous lines was obtained.Cytological analysis of the mutant and functional analysis of the candidate gene Osfbox have been conducted.The main results are described as follows:
1.The transformation positive ratio of 9024 transformants was checked by using GAL4/VP16 fragment in T-DNA as a PCR marker,7862(87.1%) showed to be PCR positive.
2.Employing the TAIL-PCR method,I independently isolated 5100 FSTs,2609 (51.2%) out of which can be well mapped onto the dee genome.Combined with the FSTs isolated by colleagues,there are totally 30577 FSTs in our group, including 51.5%can be mapped.
3.T-DNA insertions were biased towards large chromosomes,not only in the absolute number of insertions but also in the relative density,and the insertion density is highly correlated with the chromosome size(r= 0.74,P＜0.01).Within chromosomes,the insertions occurred more densely in the distal ends,and less densely in the centromeric regions and the insertion number is significantly correlated with the cDNA number in corresponding region(r=0.70,P＜0.01); T-DNA insertions strongly disfavored transposable element(TE)-related sequences(SR=—34.0) and favored genic sequences(SR=15.55).In rice genes, strong bias toward the 5' upstream(SR=12.8)and Y downstream regions(SR=10.3) of the genes were detected respectively,but the coding sequence region is observed to be disfavored(SR=—13.2).In the coding region,no insertional bias between intron and exon region was observed(χ~2 = 2.81,P = 0.09).
4.According to the "molecular functions" of genes,T-DNA insertions bias among the various classes of functional genes were observed(χ~2=93.5,P＜0.000001): preferentially occurred in "Antioxidant"(SR =6.07)and "Catalytic"(SR=3.79), but preferentially not occurred in "Nutrient reservoir","Enzyme regulator", "Transcription regulator" and "Ligand binding and cartier"(SR between -2.1 to -3.8).In the other 5 gene classes,T-DNA showed a random distribution pattern.
5.In ISNS bendability analysis result,the random CK showed an irregular curve. But in our ISNS,elevated bendability was observed at positions from -200 to 200 bp and the bendability peak is symmetric in this region,with the highest points at both -10 and 10 bp from the insertion sites.There was an abrupt drop in the bendability value at the insertion sites.A similar result was also observed in ISNS from other 3 research groups.
6.The analysis of ISNS GC content indicated that GC content is not a necessary factor for T-DNA insertion site.But The GC skew and TA skew calculation exhibited several features in both ISNS from our lab and 3 other groups:The GC and TA skews appeared to be inversely correlated with r＞-0.92;The two curves crossed each other at the insertion sites(point 0) at which both GC and TA skews were equal to 0;From the insertion site to 800-bp upstream,the GC skew was positive and reached a plateau in approximately the region from -300 to -100 bp, whereas the TA skew was negative and formed a valley in a similar position in approximately the region from -300 to -100 bp.The reverse was the case from the insertion sites to 800 bp downstream,whereas no such features were observed in the 2000 random sequences.
7.In our T-DNA insertional population,T-DNA inserted in one loci as a "tandem repeats" was widely detected.The ratio of "direct repeat","5' end junction inverse repeat" and "3' end junction inverse repeat" can be up to 19.9%,10.5% and 25.4%respectively.40.9%are detected to have at least one tandem repeat. The introduction of vector backbone sequence into the rice genome are prominent as well,up to 49.6%and 29.7%of the transformants habor a backbone sequence in left and right border respectively.
8.Reverse genetics study was carried out in a mutant which harboring a T-DNA insertion in the 3'UTR of an f-box gene in chromosome 6.The T-DNA hemizygous lines are semi-sterile,but both the T-DNA negative and homozygous lines are full fertile.This is similar to the semi-sterility of indica-japonica subspecies hybrids.The mutant phenotype is checked to be co-segregated with the T-DNA insertion.The candidate gene is named as Osfoox.
9.The ID of Osfoox in TIGR is LOC_Os06g06050 with a length of 4441bp.The full-length eDNA information in KOME(AK065478) showed that:3723bp in length,containes 4 exons and 3 introns,encoding a 720AA protein.A potential F-box domain in 15-63 AA and Cysteine-eontaining LRR profile domain in 390-469AA were detected by Plantsp.
10.Cytological analysis showed that tapetum degeneration retardation happened in tetrad stage,and the semi-sterility of T-DNA hemizygous lines are a consequence of both the undeveloped microspore and megaspore,which can pass the meiosis stage but are blocked in the following mitosis stage.
11.By comparing the Osfbox gene sequence of 10 typical rice indica or japonica varieties,we found 18 base variations between the 2 subspecies,including 12 occurred in the 648bp upstream region from the transcription start site.In gene coding region,only 2 deletions were detected,resulting in a 3 tandem Glu and 1 Gly deletion respectively.
12.Microarray based expression profile and RT-PCR data showed that Osfbox is a constitutively expressed gene.A higher expression level was detected in anther and pistil.In young panicle development stage 3-8,Osfbox expression level reaches the peak in stage 4 and then decreased.In situ hybridization result also showed that Osfbox is strongly expressed until tetrad formatted,but after the vacuolated pollen stage,no expression was detected in pollens.
13.The Osfbox protein is specifically localized in cell nuclear.
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