水稻窄卷叶基因NRL1的克隆与功能验证&水稻杂种弱势的遗传机理研究
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摘要
水稻叶片是植物进行光合作用的主要营养器官，也是理想株型的重要组成部分，因此水稻叶型已经成为了分子遗传和育种学家研究的热点。为了进一步了解水稻叶发育的调控机制，本研究从日本晴和武运粳7号的EMS诱变中成功分离到三个等位的窄卷叶突变体nrl1-1、nrl1-2和nrl1-3，它们都表现出了矮化、窄叶微卷、籽粒细长，根系变短等一系列形态上的变化。切片观察表明NRL1能够控制一系列细胞和组织的发育，叶片的泡状细胞、叶脉、茎秆厚壁细胞和薄壁细胞都在nrl1内发生了大小或数量上的变化，其中泡状细胞的缩小，能够造成nrl1叶片向内卷曲，而叶脉数目的下降也导致了nrl1叶片的窄化。同时突变体内茎秆的纤维素、半纤维素和木质素含量都有所提高，因此推测NRL1很可能涉及到了它们的生物合成。
通过图位克隆法最终将NRL1定位在第12染色体8.7kb的范围内，该区间只有一个开放阅读框。测序后发现，三个等位突变体都在该基因的编码序列上发生了碱基变化，通过构建的NRL1功能互补和过表达载体进行了转基因试验，结果表明，突变体转互补载体和过表达载体的植株表型都得到了恢复，但野生型日本晴转互补和过表达载体的植株并没有增加叶片的宽度和卷曲度，表明NRL1没有剂量效应。NRL1在组织中表现为组成型表达特征，其在所测试的根、茎、叶、鞘、穗上都表达，在突变体中的表达量有所下降，在互补转基因植株中表达差异不明显，但在过表达植株中增加显著。序列分析表明NRL1编码了一个类纤维素合成蛋白D4（OsCslD4），由1215个氨基酸组成，共包含了两个外显子和一个内含子。与其它Csl家族一样，NRL1同样包含了结合核苷酸糖和催化纤维素合成的D,D,D,QXXRW保守基序和八个跨膜区。
杂种弱势是植物中普遍存在的一种合子后生殖隔离现象，对杂种弱势控制基因的研究将为进一步了解生殖隔离的分子作用机理和生产应用打下理论基础。本研究发现秘鲁的品种Jamaica和常规粳稻品种杂交产生的F1均为弱势，植株表现为矮化、单分蘖、短根等特征。杂种F1种子在胚胎发育和种子萌发过程中都正常，但生长势在萌发后受到抑制，并且根系受到的抑制比苗早。其在常规大田管理条件下无法存活，但如果避免生存竞争，并给予良好的生长条件，杂种弱势植株可以存活到成熟，并可少量结实。遗传分析表明，该杂种弱势受显性互补基因Hwc1和Hwc2控制，其中Jamaica的基因型为Hwc1Hwc1hwc2hwc2，粳稻的基因型为hwc1hwc1Hwc2Hwc2，而籼稻的基因型则为hwc1hwc1hwc2hwc2，当Hwc1和Hwc2位点同时显性纯合或显性杂合时表现为弱势，而当两位点有一个位点隐性纯合或两位点都隐性纯合时就表现正常。
通过上清液PCR快速检测和测交验证重组个体，Hwc1最终被定位在第1染色体22.9kb的范围内，该区间只有一个开放阅读框，Hwc2也被确定在第4染色体24.6kb的区间内，该区段共有4个开放阅读框。目前已经构建好了相应候选基因的功能互补载体，转基因试验正在进行中。为了在生产上应用杂种弱势基因，我们将Jamaica所特有的Hwc1导入到抗虫BT、抗除草剂农达和Basta的转基因植株中，以建立“无外源基因逃逸”的转基因技术，现已经获得了抗农达、Basta和BT与Hwc1的基因聚合株。同时也将Hwc1导入到保持系武运粳7号和恢复系C堡中，现已分别获得了6代和7代的近等基因系，为Hwc1基因在杂交水稻中的利用奠定了基础。
Leaf is a main organ of photosynthesis and contributes to ideal plant type in rice. Therefore, leafshape has become one of hot spots by molecular geneticist and breeder. In order to furtherunderstanding of the regulatory mechanism of leaf development, we isolated three allelic mutants ofnrl1, nrl1-1and nrl1-2from an M2population of the japonica rice cultivar Nipponbare, nrl1-3from thejaponica rice cultivar Wuyunjing7after EMS mutagenesis. Three mutants showed phenotypes ofdwarfism with different degree, reduced leaf width, semi-rolled leaves and slim grain. Microscopicanalysis indicated that the NRL1involved in development of different cells and tissues, i.e. the bulliformcells, longitudinal veins, sclerenchyma cells and parenchyma cells were changed in size or number.Among them, smaller bulliform cells may explain the semi-rolled leaves and fewer leaf veins result inthe narrowed leaves. The content of cellulose, hemicellulose and lignin also increased in mutants,suggesting NRL1likely affect their biosynthesis.
The NRL1gene was fine mapped to an interval of8.71kb, where one open reading frame (ORF)located on chromosome12via map-based cloning strategy. To define the mutation locus, the DNAfragments of wild-type and three allelic mutants were amplified and sequenced. Single base-pairsubstitutions were found at different loci in each of the three allelic mutants. To confirm the biologicalfunction of the NRL1gene, we constructed complementation vector and over-expression vector. Thephenotype of mutant was completely suppressed while Nipponbare not changed in complementation testand over-expression analysis which confirming NRL1without dosage effect. NRL1exhibitedconstitutive expression at transcription level at the heading stage, and decreased in the mutants, alteredlittle in complementary transgenic plants and increased significantly in over-expression plants.Sequence analysis of cDNA revealed that the NRL1gene is composed of two exons and one intron,which encodes a protein of1215amino acids in the KOME database. BLAST analysis indicated that theNRL1protein is the cellulose synthase-like protein D4(OsCslD4), which contains the D, D, D,QXXRW motif and eight trans-membrane segments characterized in Csl family.
As the phenomenon of postzygotic reproduction isolation, hybrid weakness is common in plants.Genetic study of hybrid weakness will contribute to understanding the molecular mechanism ofreproduction isolation and its application. In this study, the F1plants from the cross between Peruvianvariety Jamaica and japonica varieties exhibited phenotypes of dwarfism, single tiller and notablyshortened roots. The F1seeds were normal in embryonic development and germination, but hybridweakness exhibited weak growth potential after germination, while growth was inhibited in root earlierthan in shoot. The plants will die under conventional field management, but flowering and seed settingwas achieved under favorable conditions without survival competition. Genetic analysis indicatedhybrid weakness was controlled by a pair of dominant complementary genes. The genotype of Jamaica,japonica and indica rice is Hwc1Hwc1hwc2hwc2, hwc1hwc1Hwc2Hwc2and hwc1hwc1hwc2hwc2,respectively. When the locus of Hwc1and Hwc2are dominant homozygous or dominant heterozygous,the plant appears hybrid weakness and it is normal if one or two loci is recessive homozygous.
The hybrid weakness gene Hwc1was fine mapped into a region of22.9kb on chromosome1,which had only one ORF by PCR rapid screening and test cross of recombinant. Hwc2was also finemapped in24.6kb on chromosome4, where total four genes were predicted. Complementation vectorsfor both genes have constructed and transgenic experiment is being carried out. In order to apply thehybrid weakness gene in rice production, we introduced Hwc1into a transgenic plant with Bt, Roundupand Bar gene. Now, herbicide and pest resistant plants with Hwc1have obtained by gene pyramiding toestablish a new technology of preventing the exogenous gene escape. The Hwc1also has beenintroduced into maintainer line Wuyunjing7and restorer line C Bao by backcrosses and have gainedNILs of BC6and BC7respectively, which provides basis for production apllication in hybrid rice.
通过图位克隆法最终将NRL1定位在第12染色体8.7kb的范围内，该区间只有一个开放阅读框。测序后发现，三个等位突变体都在该基因的编码序列上发生了碱基变化，通过构建的NRL1功能互补和过表达载体进行了转基因试验，结果表明，突变体转互补载体和过表达载体的植株表型都得到了恢复，但野生型日本晴转互补和过表达载体的植株并没有增加叶片的宽度和卷曲度，表明NRL1没有剂量效应。NRL1在组织中表现为组成型表达特征，其在所测试的根、茎、叶、鞘、穗上都表达，在突变体中的表达量有所下降，在互补转基因植株中表达差异不明显，但在过表达植株中增加显著。序列分析表明NRL1编码了一个类纤维素合成蛋白D4（OsCslD4），由1215个氨基酸组成，共包含了两个外显子和一个内含子。与其它Csl家族一样，NRL1同样包含了结合核苷酸糖和催化纤维素合成的D,D,D,QXXRW保守基序和八个跨膜区。
杂种弱势是植物中普遍存在的一种合子后生殖隔离现象，对杂种弱势控制基因的研究将为进一步了解生殖隔离的分子作用机理和生产应用打下理论基础。本研究发现秘鲁的品种Jamaica和常规粳稻品种杂交产生的F1均为弱势，植株表现为矮化、单分蘖、短根等特征。杂种F1种子在胚胎发育和种子萌发过程中都正常，但生长势在萌发后受到抑制，并且根系受到的抑制比苗早。其在常规大田管理条件下无法存活，但如果避免生存竞争，并给予良好的生长条件，杂种弱势植株可以存活到成熟，并可少量结实。遗传分析表明，该杂种弱势受显性互补基因Hwc1和Hwc2控制，其中Jamaica的基因型为Hwc1Hwc1hwc2hwc2，粳稻的基因型为hwc1hwc1Hwc2Hwc2，而籼稻的基因型则为hwc1hwc1hwc2hwc2，当Hwc1和Hwc2位点同时显性纯合或显性杂合时表现为弱势，而当两位点有一个位点隐性纯合或两位点都隐性纯合时就表现正常。
通过上清液PCR快速检测和测交验证重组个体，Hwc1最终被定位在第1染色体22.9kb的范围内，该区间只有一个开放阅读框，Hwc2也被确定在第4染色体24.6kb的区间内，该区段共有4个开放阅读框。目前已经构建好了相应候选基因的功能互补载体，转基因试验正在进行中。为了在生产上应用杂种弱势基因，我们将Jamaica所特有的Hwc1导入到抗虫BT、抗除草剂农达和Basta的转基因植株中，以建立“无外源基因逃逸”的转基因技术，现已经获得了抗农达、Basta和BT与Hwc1的基因聚合株。同时也将Hwc1导入到保持系武运粳7号和恢复系C堡中，现已分别获得了6代和7代的近等基因系，为Hwc1基因在杂交水稻中的利用奠定了基础。
Leaf is a main organ of photosynthesis and contributes to ideal plant type in rice. Therefore, leafshape has become one of hot spots by molecular geneticist and breeder. In order to furtherunderstanding of the regulatory mechanism of leaf development, we isolated three allelic mutants ofnrl1, nrl1-1and nrl1-2from an M2population of the japonica rice cultivar Nipponbare, nrl1-3from thejaponica rice cultivar Wuyunjing7after EMS mutagenesis. Three mutants showed phenotypes ofdwarfism with different degree, reduced leaf width, semi-rolled leaves and slim grain. Microscopicanalysis indicated that the NRL1involved in development of different cells and tissues, i.e. the bulliformcells, longitudinal veins, sclerenchyma cells and parenchyma cells were changed in size or number.Among them, smaller bulliform cells may explain the semi-rolled leaves and fewer leaf veins result inthe narrowed leaves. The content of cellulose, hemicellulose and lignin also increased in mutants,suggesting NRL1likely affect their biosynthesis.
The NRL1gene was fine mapped to an interval of8.71kb, where one open reading frame (ORF)located on chromosome12via map-based cloning strategy. To define the mutation locus, the DNAfragments of wild-type and three allelic mutants were amplified and sequenced. Single base-pairsubstitutions were found at different loci in each of the three allelic mutants. To confirm the biologicalfunction of the NRL1gene, we constructed complementation vector and over-expression vector. Thephenotype of mutant was completely suppressed while Nipponbare not changed in complementation testand over-expression analysis which confirming NRL1without dosage effect. NRL1exhibitedconstitutive expression at transcription level at the heading stage, and decreased in the mutants, alteredlittle in complementary transgenic plants and increased significantly in over-expression plants.Sequence analysis of cDNA revealed that the NRL1gene is composed of two exons and one intron,which encodes a protein of1215amino acids in the KOME database. BLAST analysis indicated that theNRL1protein is the cellulose synthase-like protein D4(OsCslD4), which contains the D, D, D,QXXRW motif and eight trans-membrane segments characterized in Csl family.
As the phenomenon of postzygotic reproduction isolation, hybrid weakness is common in plants.Genetic study of hybrid weakness will contribute to understanding the molecular mechanism ofreproduction isolation and its application. In this study, the F1plants from the cross between Peruvianvariety Jamaica and japonica varieties exhibited phenotypes of dwarfism, single tiller and notablyshortened roots. The F1seeds were normal in embryonic development and germination, but hybridweakness exhibited weak growth potential after germination, while growth was inhibited in root earlierthan in shoot. The plants will die under conventional field management, but flowering and seed settingwas achieved under favorable conditions without survival competition. Genetic analysis indicatedhybrid weakness was controlled by a pair of dominant complementary genes. The genotype of Jamaica,japonica and indica rice is Hwc1Hwc1hwc2hwc2, hwc1hwc1Hwc2Hwc2and hwc1hwc1hwc2hwc2,respectively. When the locus of Hwc1and Hwc2are dominant homozygous or dominant heterozygous,the plant appears hybrid weakness and it is normal if one or two loci is recessive homozygous.
The hybrid weakness gene Hwc1was fine mapped into a region of22.9kb on chromosome1,which had only one ORF by PCR rapid screening and test cross of recombinant. Hwc2was also finemapped in24.6kb on chromosome4, where total four genes were predicted. Complementation vectorsfor both genes have constructed and transgenic experiment is being carried out. In order to apply thehybrid weakness gene in rice production, we introduced Hwc1into a transgenic plant with Bt, Roundupand Bar gene. Now, herbicide and pest resistant plants with Hwc1have obtained by gene pyramiding toestablish a new technology of preventing the exogenous gene escape. The Hwc1also has beenintroduced into maintainer line Wuyunjing7and restorer line C Bao by backcrosses and have gainedNILs of BC6and BC7respectively, which provides basis for production apllication in hybrid rice.
引文
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