水稻HD-ZIP转录因子Oshox4，Oshox6和IPT基因在水稻中的表达及其抗旱功能研究
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摘要
水稻是世界主要粮食作物之一，各种非生物逆境如干早、高低温以及高盐等对作物的生长和发育有着非常重要的影响，从而导致产量下降。在许多地区，干旱作为最普遍的非生物逆境形式，己成为农业发展的瓶颈。水稻是我国的主要粮食作物，而且需水量巨大。因此进行水稻抗旱性研究，减少农业生产用水，提高水稻的抗旱性并培育出抗旱性品种显得尤为重要。
本研究以获得水稻抗旱遗传改良基因为目的，通过在水稻中过量表达对抗旱调控起核心作用的转录因子和相关抗旱基因，对其在大田和温室中的抗旱性进行系统研究分析。初步确定转录因子基因（Oshox4和Oshox6）和IPT(iso pentenyl transferases)基因在水稻抗旱遗传改良中具有潜在价值，最后对以上3个基因的抗旱机制分别进行了较为深入研究。主要结果如下：
1．构建了35s::Oshox4PCAMBIA1300过表达载体，以农杆菌介导法转化抗旱胁迫敏感型水稻品种（IR64）,得到了54株阳性转基因植株。通过southern杂交分析，从54个转化植株中鉴定出16个单拷贝株系。对以上16个单拷贝的转基因株系的T1代进行了生理表型观察和研究。
2．选择了2个单拷贝株系在开花期对其进行干旱胁迫并测定了干旱指标，确定了Oshox4基因干旱胁迫下的功能。观察6个单拷贝Oshox4转化株系T2代的大田农艺性状，相对于野生型株系，6个转基因株系分蘖数增多、穗数明显增加、株高变矮、茎间缩短、开花期延后且子粒不饱满。
3.通过TAIL-PCR找出了目的基因在染色体的插入位点，并在此基础上设计特异引物从T1代鉴定出纯合系。通过表型观察发现，同一株系的纯合和杂合植株之间有明显表型差异。
4．构建了Oslea：：Oshox6PCAMBIA1300干旱胁迫诱导型表达载体，以农杆菌介导法转化抗旱胁迫敏感型水稻品种（IR64），得到86个潮霉素抗性再生植株。通过PCR和Southern杂交分析确定了目的基因的整合以及拷贝数。对3个Oshox6单拷贝转化株的T2代在PVC管进行干旱胁迫，3个转基因株系的根长和根直径均显著高于野生型。在大田里干旱胁迫中，转基因株系的抗旱性也明显优先于野生型。
5.克隆到Oshox6基因启动子，并连接到Gus基因，构建了Oshox6pro：：Gus PCAMBIA1300载体，以农杆菌介导法转化转化到水稻（IR64）。通过PCR确定了阳性植株，并对阳性株的根、茎和叶片进行Gus染色，结果表明Oshox6启动子在水稻根的木质部特异性表达。
6．构建了SARK：：IPT PCAMBIA1380逆境诱导表达载体，以农杆菌介导法转化转化到水稻（IR64），得到了256个潮霉素抗性再生植株。通过PCR和Southern杂交分析确定了目的基因在基因组的整合以及拷贝数。根据southern杂交结果选取了10个单拷贝的IPT阳性植株进行农艺性状观察。对两个转基因株系进行干旱胁迫，结果表明，在干旱胁迫条件下IPT转基因水稻的叶绿素和细胞分裂素含量显著高于野生型。
Rice is one of the world’s most important food crops, various of biological stress such as drought,cold, high temperature and salinity is significantly impact on plant growth and development. Resulting isdecline yield. Drought is general form of biological stress, and already becomes a big problem foragricultural developing in many places. rice is major crops in our country and it needs huge water duringthe life cycle.so to studying drought tolerance of rice, reducing the water for agricultural production,improve the drought tolerance of rice and cultivate drought resistance varieties is particularly important
In this study, to obtain a gene that enhanced rice drought genetic improvement as purpose,through overstressing of transcription factor and association with drought tolerance gene in rice, and ingreen house and field systematically analyze for the transgenic plant.Initially identified transcriptionfactor gene (Oshox4and Oshox6) and IPT (isopentenyltransferases) gene have a potential value ofgenetic improvement of rice drought tolerance, at the end for the three gene we did a more in depthstudy. main results are as below:
1.35S::Oshox4PCAMBIA vector were constructed, through Agro bacteriumtumefactions-mediated the gene transfer to drought stress sensitive variety IR64, and54positive plantswere obtained. by southern blotting analyze from54transgenic events19individual events were singlecopy,were confirm individual plants was positive, for the19individual single copy events and wild typeplant carried out phenotype observation and analyze.
2. Selected two transgenic single copy events exposed to progressive soil drying at flowering stageand measured some drought index, to confirm Oshox4gene function in the drought stress condition. Andsame time we carried out phenotype observation for6single copy T2generation plants, found thatcompare to wild type all the transgenic events number of tiller and panicle were increased, but plantheight and internodes were shorter than wild type, flowering date is delayed and grains were unfilled.
3. Using TAIL-PCR method found gene insert position in the chromosome and depend on geneinsert position design specific primer for the each line determine homozygous plants form T1generation.through phenotype observation found that in the same line between homozygous andheterozygous also have a obvious difference.
4. Constructed PCAMBIA1300Oslea：：Oshox6drought induce able vector, through agrobacterium tumefactions-mediated transform to drought sensitive rice variety IR64,obtained86hygromycin resistance plants. Using PCR and Southern blot analyze confirm gene integration and copynumber. The three single copy T2generation plants were exposing drought stressed in the PVC tube, andobserved that three transgenic plants root length and root diameter were obviously higher than wildtype.In the field drought stress condition,transgenic plant’s drought tolerance is better than wild type.
5. Oshox6gene natural promoter was cloned and ligated with Gus gene,made a Oshox6promoter：：Gus construct, Through Agro bacterium tumefactions-mediated transform to rice variety,using PCR confirm gene integration and processing Gus staining to the transgenic plant root, shoot leafand anther, we found that Oshox6promoter was specific expression in root endodermis.
6. SARK：：IPT PCAMBIA1380drought induce able vector were constructed, through Agrobacterium tumefacients-mediated transform to rice variety IR64,obtained256hygromycin resistanceplant,by the PCR and Southern blot analyze to confirm gene integration and copy number.depend onSouthern blot analyze selected10single copy plants were selected for the phenotype observation. Andexposed to drought stress for the two transgenic plants, the result showed that in drought stress conditionIPT transgenic plants chlorophyll content and cytokine content were obviously higher than wild type.
本研究以获得水稻抗旱遗传改良基因为目的，通过在水稻中过量表达对抗旱调控起核心作用的转录因子和相关抗旱基因，对其在大田和温室中的抗旱性进行系统研究分析。初步确定转录因子基因（Oshox4和Oshox6）和IPT(iso pentenyl transferases)基因在水稻抗旱遗传改良中具有潜在价值，最后对以上3个基因的抗旱机制分别进行了较为深入研究。主要结果如下：
1．构建了35s::Oshox4PCAMBIA1300过表达载体，以农杆菌介导法转化抗旱胁迫敏感型水稻品种（IR64）,得到了54株阳性转基因植株。通过southern杂交分析，从54个转化植株中鉴定出16个单拷贝株系。对以上16个单拷贝的转基因株系的T1代进行了生理表型观察和研究。
2．选择了2个单拷贝株系在开花期对其进行干旱胁迫并测定了干旱指标，确定了Oshox4基因干旱胁迫下的功能。观察6个单拷贝Oshox4转化株系T2代的大田农艺性状，相对于野生型株系，6个转基因株系分蘖数增多、穗数明显增加、株高变矮、茎间缩短、开花期延后且子粒不饱满。
3.通过TAIL-PCR找出了目的基因在染色体的插入位点，并在此基础上设计特异引物从T1代鉴定出纯合系。通过表型观察发现，同一株系的纯合和杂合植株之间有明显表型差异。
4．构建了Oslea：：Oshox6PCAMBIA1300干旱胁迫诱导型表达载体，以农杆菌介导法转化抗旱胁迫敏感型水稻品种（IR64），得到86个潮霉素抗性再生植株。通过PCR和Southern杂交分析确定了目的基因的整合以及拷贝数。对3个Oshox6单拷贝转化株的T2代在PVC管进行干旱胁迫，3个转基因株系的根长和根直径均显著高于野生型。在大田里干旱胁迫中，转基因株系的抗旱性也明显优先于野生型。
5.克隆到Oshox6基因启动子，并连接到Gus基因，构建了Oshox6pro：：Gus PCAMBIA1300载体，以农杆菌介导法转化转化到水稻（IR64）。通过PCR确定了阳性植株，并对阳性株的根、茎和叶片进行Gus染色，结果表明Oshox6启动子在水稻根的木质部特异性表达。
6．构建了SARK：：IPT PCAMBIA1380逆境诱导表达载体，以农杆菌介导法转化转化到水稻（IR64），得到了256个潮霉素抗性再生植株。通过PCR和Southern杂交分析确定了目的基因在基因组的整合以及拷贝数。根据southern杂交结果选取了10个单拷贝的IPT阳性植株进行农艺性状观察。对两个转基因株系进行干旱胁迫，结果表明，在干旱胁迫条件下IPT转基因水稻的叶绿素和细胞分裂素含量显著高于野生型。
Rice is one of the world’s most important food crops, various of biological stress such as drought,cold, high temperature and salinity is significantly impact on plant growth and development. Resulting isdecline yield. Drought is general form of biological stress, and already becomes a big problem foragricultural developing in many places. rice is major crops in our country and it needs huge water duringthe life cycle.so to studying drought tolerance of rice, reducing the water for agricultural production,improve the drought tolerance of rice and cultivate drought resistance varieties is particularly important
In this study, to obtain a gene that enhanced rice drought genetic improvement as purpose,through overstressing of transcription factor and association with drought tolerance gene in rice, and ingreen house and field systematically analyze for the transgenic plant.Initially identified transcriptionfactor gene (Oshox4and Oshox6) and IPT (isopentenyltransferases) gene have a potential value ofgenetic improvement of rice drought tolerance, at the end for the three gene we did a more in depthstudy. main results are as below:
1.35S::Oshox4PCAMBIA vector were constructed, through Agro bacteriumtumefactions-mediated the gene transfer to drought stress sensitive variety IR64, and54positive plantswere obtained. by southern blotting analyze from54transgenic events19individual events were singlecopy,were confirm individual plants was positive, for the19individual single copy events and wild typeplant carried out phenotype observation and analyze.
2. Selected two transgenic single copy events exposed to progressive soil drying at flowering stageand measured some drought index, to confirm Oshox4gene function in the drought stress condition. Andsame time we carried out phenotype observation for6single copy T2generation plants, found thatcompare to wild type all the transgenic events number of tiller and panicle were increased, but plantheight and internodes were shorter than wild type, flowering date is delayed and grains were unfilled.
3. Using TAIL-PCR method found gene insert position in the chromosome and depend on geneinsert position design specific primer for the each line determine homozygous plants form T1generation.through phenotype observation found that in the same line between homozygous andheterozygous also have a obvious difference.
4. Constructed PCAMBIA1300Oslea：：Oshox6drought induce able vector, through agrobacterium tumefactions-mediated transform to drought sensitive rice variety IR64,obtained86hygromycin resistance plants. Using PCR and Southern blot analyze confirm gene integration and copynumber. The three single copy T2generation plants were exposing drought stressed in the PVC tube, andobserved that three transgenic plants root length and root diameter were obviously higher than wildtype.In the field drought stress condition,transgenic plant’s drought tolerance is better than wild type.
5. Oshox6gene natural promoter was cloned and ligated with Gus gene,made a Oshox6promoter：：Gus construct, Through Agro bacterium tumefactions-mediated transform to rice variety,using PCR confirm gene integration and processing Gus staining to the transgenic plant root, shoot leafand anther, we found that Oshox6promoter was specific expression in root endodermis.
6. SARK：：IPT PCAMBIA1380drought induce able vector were constructed, through Agrobacterium tumefacients-mediated transform to rice variety IR64,obtained256hygromycin resistanceplant,by the PCR and Southern blot analyze to confirm gene integration and copy number.depend onSouthern blot analyze selected10single copy plants were selected for the phenotype observation. Andexposed to drought stress for the two transgenic plants, the result showed that in drought stress conditionIPT transgenic plants chlorophyll content and cytokine content were obviously higher than wild type.
引文
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