水稻（Oryza sativa L.）光呼吸突变体线粒体丝氨酸羟甲基转移酶基因克隆及功能分析
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摘要
水稻(Oryza sativa L.)作为世界上最重要的粮食作物之一,基因组全序列的获得和大规模突变体库的建立,为水稻功能基因组学开展创造了良好条件。目前,许多涉及重要农艺形状的突变体已被鉴定,并且基因也陆续被克隆,这为了解水稻农艺形状形成的遗传基础做出了贡献。叶绿素缺失突变由于和光合作用密切相关,因此对这一类型农艺性状形成机制的研究有重要意义。本研究在获得大量水稻叶绿素缺失突变体基础上,开展了基因克隆和突变体鉴定工作,得到了一个参与光呼吸途径的基因(OsmSHMT),现将主要研究内容概括如下：
1突变体表型鉴定
通过对生长于培养室内的突变体和野生型观察发现突变体表型一开始并不明显,播种后15天叶绿色缺失表型才逐渐得以区分。在20天之前,突变体生长较野生型快,幼苗纤细,由于无法支撑进一步伸展的叶片,植株逐渐弯曲。
2突变基因克隆
利用GUS染色初步确定了突变表型可能与T-DNA插入有关,经过PCR-walking分离了突变体基因组T-DNA插入位点侧翼序列,通过比对后发现T-DNA插入水稻线粒体丝氨酸羟甲基转移酶基因(OsmSHMT),经PCI共分离验证,确定了被标记基因与突变表型共分离。最后,经互补实验证实OsmSHMT基因突变造成叶绿素缺失表型。
3 OsmSHMT基因表达模式
利用增强子捕获载体功能,经GUS染色探讨了OsmSHMT基因在各组织器官表达模式,结果表明该基因主要在叶片等具有光合功能的器官和部位表达,除叶片外,在根部也有强烈表达,而花器官内表达很弱。
4 OsmSHMT.突变体在高CO2浓度下回复为野生表型
由基因生物信息学分析认为OsmSHMT基因可能参与光呼吸途径,而光呼吸突变体都能在高CO2浓度下正常生长,不表现突变表型,为验证这一点,观察了OsmSHMT突变体在高CO2浓度环境中生长状况,结果证实OsmSHMT突变体属于光呼吸突变体。
5 OsmSHMT基因原核表达
在构建OsmSHMT-pET28a基础上,将其转入寄主菌BL21中,实现了OsmSHMT基因原核表达。
6 OsmSHMT基因突变对光合作用影响
利用叶绿素荧光技术,通过对突变体和野生型荧光慢诱导曲线测定发现,突变体荧光淬灭能力明显下降,高光强下尤其明显。通过观察qP和qN变化发现,qN在低光强下即表现出与野生型明显差异,而qP则在高光强下差异明显。经分析后认为,突变体qN降低可能与类囊体膜透性增加有关,qP可能受卡尔文循环影响而降低。7叶绿体结构及类囊体蛋白复合物变化
叶绿体电镜切片表明,与野生型相比,突变体叶肉细胞内叶绿体数量减少,体积变小,基粒分布明显不均,基质类囊体破坏明显,嗜锇体增多。叶绿体类囊体膜蛋白复合物温和胶电泳结果表明,相比野生型,突变体光系统主要大分子复合物带型上移,这可能是由于受到氧化伤害后大分子复合物结构松散体积增大所致。
综上所述水稻线粒体丝氨酸羟甲基转移酶基因(OsmSHMT)突变导致光呼吸途径受损,进而突变体卡尔文循环受到影响,光能不能被有效利用,过剩光能导致叶绿体活性氧升高,造成叶绿素、叶绿体膜系统及类囊体蛋白复合物被光氧化,叶绿体数量减少体积变小,最终,表型上表现为突变体叶绿素缺失性状。
The sequencing of rice genome, a important crop, and constructing of T-DNA insertion mutant pools have provided good conditions for rice functional genomics. Many important agricultural character mutants have been isolated so far, in which some genes causing mutant phenotypes have also been cloned. These works are beneficial for learning the genetics of agricultural characters. For closely related to photosynthesis, the studying of chlorophyll deficient mutations is significance. In this study, after obtaining lots of chlorophyll deficient mutants, a gene relation to photorespiration was cloned and the functions were studied. The most studying contents are listed as follows: 1 Identification of mutant phenotype
When planted in culture room, the mutant phenotype was difficult to discern from wild at first, but after 15 days since sowing, the mutant phenotype appeared gradually. Before dying, the mutants grew faster than wild, but the seedlings were so slender that it couldn't sustain the weight of leafs and began to be curved. 2 Cloning of the mutated gene
It was supposed that the mutant phenotype was relation to T-DNA by GUS staining. Using PCR-walking technique, the flanking sequence of T-DNA left border was isolated easily. After BLAST, the T-DNA was proven to be inserted into a gene, predicted coding rice mitochondria serine hydroxymethyltransferase (SHMT). It was certain that the tagged gene co-segregated with mutant phenotype by PCR method. The normal gene in wild was amplified and transformed to mutant, and after regeneration, the mutant phenotype disappeared. The complementary experiment proven the cloned gene is responsible for the chlorophyll deficient mutant phenotype.
3 The expression patterns of OsmSHMT Using enhancer trap vector which can indicate expression patterns of tagged gene, the different parts were examined with GUS assay. The leaf blade and root were stained strongly, but the flower was barely.
4 The mutant phenotype was restored in high CO2 concentration
From bioinformatic analysis, the OsmSHMT gene takes part in photorespiration process. For proving the result, the OsmSHMT mutant was cultured in high CO2 concentration, after 2 weeks, no any observable mutant phenotype could be discerned from wild.
5 OsmSHMT gene prokaryotic expression
After transformation of OsmSHMT-pET28a expression vector into E.coli strain BL21(DE3), high yield of recombinant protein was achieved.
6 The changes of photosynthesis in mutant
For determining the changes of photosynthesis, the chlorophyll fluorescence curve was examined, and it observed that the chlorophyll fluorescence quenching was influenced greatly in mutant, especially in high light intensity. In low light intensity, comparing qP with qN, it was evident that the qN was declined more seriously. It might be caused by the enhanced membrane permeability. In high light intensity, the qP was also declined serious, it might be caused by damaged Calvin cycle.
7 The changes of chloroplast ultrastructure and protein complexes on thylakoid
Compared with wild type, the quantity and volume of chloroplasts in mutant mesophyll were decrease. In chloroplast of wild, the granum thylakoids were distributed more regular than mutant. It might be caused by the stroma thylakoid damage. As for the changes of protein complexes on thylakoid, from BN-gel, it was observed that most strips of mutant moved slower than wild. It might be caused by the loose structure of injured protein complexes.
In conclusion, the mutation of OsmSHMT damages photorespiration, and because the Calvin cycle needs photorespiration to sustain, the injured Calvin cycle leads to declined light energy converting and the excess light energy promotes reactive oxygen species(ROS) producing, then the membrane, the protein complexes and the chlorophyll were all injured by ROS, at last, the chlorophyll deficient mutant phenotype appears.
1突变体表型鉴定
通过对生长于培养室内的突变体和野生型观察发现突变体表型一开始并不明显,播种后15天叶绿色缺失表型才逐渐得以区分。在20天之前,突变体生长较野生型快,幼苗纤细,由于无法支撑进一步伸展的叶片,植株逐渐弯曲。
2突变基因克隆
利用GUS染色初步确定了突变表型可能与T-DNA插入有关,经过PCR-walking分离了突变体基因组T-DNA插入位点侧翼序列,通过比对后发现T-DNA插入水稻线粒体丝氨酸羟甲基转移酶基因(OsmSHMT),经PCI共分离验证,确定了被标记基因与突变表型共分离。最后,经互补实验证实OsmSHMT基因突变造成叶绿素缺失表型。
3 OsmSHMT基因表达模式
利用增强子捕获载体功能,经GUS染色探讨了OsmSHMT基因在各组织器官表达模式,结果表明该基因主要在叶片等具有光合功能的器官和部位表达,除叶片外,在根部也有强烈表达,而花器官内表达很弱。
4 OsmSHMT.突变体在高CO2浓度下回复为野生表型
由基因生物信息学分析认为OsmSHMT基因可能参与光呼吸途径,而光呼吸突变体都能在高CO2浓度下正常生长,不表现突变表型,为验证这一点,观察了OsmSHMT突变体在高CO2浓度环境中生长状况,结果证实OsmSHMT突变体属于光呼吸突变体。
5 OsmSHMT基因原核表达
在构建OsmSHMT-pET28a基础上,将其转入寄主菌BL21中,实现了OsmSHMT基因原核表达。
6 OsmSHMT基因突变对光合作用影响
利用叶绿素荧光技术,通过对突变体和野生型荧光慢诱导曲线测定发现,突变体荧光淬灭能力明显下降,高光强下尤其明显。通过观察qP和qN变化发现,qN在低光强下即表现出与野生型明显差异,而qP则在高光强下差异明显。经分析后认为,突变体qN降低可能与类囊体膜透性增加有关,qP可能受卡尔文循环影响而降低。7叶绿体结构及类囊体蛋白复合物变化
叶绿体电镜切片表明,与野生型相比,突变体叶肉细胞内叶绿体数量减少,体积变小,基粒分布明显不均,基质类囊体破坏明显,嗜锇体增多。叶绿体类囊体膜蛋白复合物温和胶电泳结果表明,相比野生型,突变体光系统主要大分子复合物带型上移,这可能是由于受到氧化伤害后大分子复合物结构松散体积增大所致。
综上所述水稻线粒体丝氨酸羟甲基转移酶基因(OsmSHMT)突变导致光呼吸途径受损,进而突变体卡尔文循环受到影响,光能不能被有效利用,过剩光能导致叶绿体活性氧升高,造成叶绿素、叶绿体膜系统及类囊体蛋白复合物被光氧化,叶绿体数量减少体积变小,最终,表型上表现为突变体叶绿素缺失性状。
The sequencing of rice genome, a important crop, and constructing of T-DNA insertion mutant pools have provided good conditions for rice functional genomics. Many important agricultural character mutants have been isolated so far, in which some genes causing mutant phenotypes have also been cloned. These works are beneficial for learning the genetics of agricultural characters. For closely related to photosynthesis, the studying of chlorophyll deficient mutations is significance. In this study, after obtaining lots of chlorophyll deficient mutants, a gene relation to photorespiration was cloned and the functions were studied. The most studying contents are listed as follows: 1 Identification of mutant phenotype
When planted in culture room, the mutant phenotype was difficult to discern from wild at first, but after 15 days since sowing, the mutant phenotype appeared gradually. Before dying, the mutants grew faster than wild, but the seedlings were so slender that it couldn't sustain the weight of leafs and began to be curved. 2 Cloning of the mutated gene
It was supposed that the mutant phenotype was relation to T-DNA by GUS staining. Using PCR-walking technique, the flanking sequence of T-DNA left border was isolated easily. After BLAST, the T-DNA was proven to be inserted into a gene, predicted coding rice mitochondria serine hydroxymethyltransferase (SHMT). It was certain that the tagged gene co-segregated with mutant phenotype by PCR method. The normal gene in wild was amplified and transformed to mutant, and after regeneration, the mutant phenotype disappeared. The complementary experiment proven the cloned gene is responsible for the chlorophyll deficient mutant phenotype.
3 The expression patterns of OsmSHMT Using enhancer trap vector which can indicate expression patterns of tagged gene, the different parts were examined with GUS assay. The leaf blade and root were stained strongly, but the flower was barely.
4 The mutant phenotype was restored in high CO2 concentration
From bioinformatic analysis, the OsmSHMT gene takes part in photorespiration process. For proving the result, the OsmSHMT mutant was cultured in high CO2 concentration, after 2 weeks, no any observable mutant phenotype could be discerned from wild.
5 OsmSHMT gene prokaryotic expression
After transformation of OsmSHMT-pET28a expression vector into E.coli strain BL21(DE3), high yield of recombinant protein was achieved.
6 The changes of photosynthesis in mutant
For determining the changes of photosynthesis, the chlorophyll fluorescence curve was examined, and it observed that the chlorophyll fluorescence quenching was influenced greatly in mutant, especially in high light intensity. In low light intensity, comparing qP with qN, it was evident that the qN was declined more seriously. It might be caused by the enhanced membrane permeability. In high light intensity, the qP was also declined serious, it might be caused by damaged Calvin cycle.
7 The changes of chloroplast ultrastructure and protein complexes on thylakoid
Compared with wild type, the quantity and volume of chloroplasts in mutant mesophyll were decrease. In chloroplast of wild, the granum thylakoids were distributed more regular than mutant. It might be caused by the stroma thylakoid damage. As for the changes of protein complexes on thylakoid, from BN-gel, it was observed that most strips of mutant moved slower than wild. It might be caused by the loose structure of injured protein complexes.
In conclusion, the mutation of OsmSHMT damages photorespiration, and because the Calvin cycle needs photorespiration to sustain, the injured Calvin cycle leads to declined light energy converting and the excess light energy promotes reactive oxygen species(ROS) producing, then the membrane, the protein complexes and the chlorophyll were all injured by ROS, at last, the chlorophyll deficient mutant phenotype appears.
引文
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