水稻Hsp17.0、DREBA5-1和RAV1-1基因功能的初步研究
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摘要
水稻是一种重要的农作物,世界半数以上的人口以大米为主食。各种非生物逆境诸如高温、干旱和高盐胁迫影响水稻的生长发育,导致水稻产量和品质受到严重的影响。利用基因工程手段和生物学技术研究水稻非生物胁迫应答过程中关键的调节基因和重要的功能基因,并阐明其分子机制,对作物遗传改良有重要意义。热激蛋白(Heat shock proteins,Hsps)是一类重要的逆境相关蛋白,能提高植物抗御逆境胁迫的能力。植物中以小分子热激蛋白(sHsps)居多AP2/EREBP(APetala2/Ethylene-responsive element binding protein)是植物所特有的一类转录因子,参与调控植物的生长发育和逆境响应。DREB和RAV是其中两个重要的亚族。为研究和分析OsHsp17.0基因和2个AP2基因(OsDREBA5-1和OsRAV1-1)的功能,本研究对过表达OsHsp17.0的转基因水稻进行了多种逆境处理；并以野生型日本晴为材料,利用半定量RT-PCR和荧光定量PCR对OsDREBA5-1和OsRAV1-1基因进行不同组织部位和高温、干旱、PEG胁迫及ABA处理的表达特征分析；构建OsDREBA5-1和OsRAV1-1基因的过表达和RNAi表达载体。主要取得了以下研究结果：
1.采用农杆菌介导法将OsHsp17.0基因的过表达载体转入水稻,从转录水平证实获得了阳性植株。对转基因植株进行盐胁迫和干旱胁迫处理后发现,OsHsp17.0-OE转基因幼苗较野生型WT出现受害症状的时间延迟,存活率提高；胁迫处理后转基因植株中脯氨酸的含量增幅大于野生型植株的,而MDA含量和离子渗漏率则相反。这些结果说明过表达OsHsp17.0提高了转基因水稻苗期对盐胁迫和干旱胁迫的耐受能力。
2.对转基因植株进行高温和低温胁迫处理后发现,OsHsp17.0-OE转基因和野生型WT幼苗均可继续生长,但二者无明显表型差异；转基因植株中脯氨酸的含量增幅大于野生型植株的,而MDA含量则相反。
3.对转基因植株进行重金属离子(镉)胁迫处理后发现,OsHsp17.0-OE转基因和野生型WT幼苗生长均受到抑制,但二者无明显表型差异。
4.两个AP2/EREBP基因OsDREBA5-1和OsRAV1-1的表达具有组织特异性。相比之下,OsDREBA5-1基因更趋向于在根部表达,而OsRAV1-1基因在叶和叶鞘表达更为强烈,推测这两个基因的功能侧重点不同。
S.OsDREBA5.1和OsRAV1-1基因在高温、PEG、NaCl和ABA处理中表达量均上调,但其表达模式有所区别。在多种逆境处理下,OsDREBA5-1基因在根部被诱导的程度要大大高于OsRAV1-1基因被诱导的程度；而在地上部分的情形恰好相反,OsRAV1-1基因被诱导的程度大大高于OsDREBA5-1基因被诱导的程度。这与这两个基因的部位表达特点相一致。OsDREBA5-1基因可能侧重于在根部参与调控对逆境的响应；而OsRAV1-1基因则更主要地在地上部分参与调控对逆境的响应。
6.构建了OsDREBA5-1和OsRAV1-1基因的过表达和RNAi干扰载体,通过农杆菌介导法获得转基因植株,并从转录水平证实获得了阳性植株。为研究这两个基因在水稻中的功能奠定了基础。
Rice is an important crop that more than half of the world people live on. Abiotic stresses such as high temperature, drought, and high salinity affect rice growth and development resulting in serious influence on rice yield and quality performance. Therefore, it is very important to find key regulatory or functional genes responsive to abiotic stresses, to reveal the molecular mechanisms of stress tolerance and to improve rice resistance by genetic engineering techniques. Heat shock proteins (Hsps) are important proteins whose expression is increased when the cells are exposed to elevated temperatures or other stresses. Increased expression of these proteins can protect the organism against stress-induced damage. Small Heat shock proteins (sHsps) are more abundant in plant. The AP2/EREBP transcription factors are known to be unique in plants, and they are involved in growth, development and environmental stress responses. DREB and RAV are two major subfamilies of AP2/EREBP. To investigate the gene function of OsHsp17.0,OsDREBA5-1 and OsRAVl-1, OsHspl7.0-overexpression rice plants have been exposed to different abiotic stresses including salt stress, drought stress, high temperature and low temperature treatment and the stress-tolerance has been analysised. Using semiquantitative RT-PCR method and real-time RT-PCR, the expression profile of OsDREBA5-1 and OsRAVl-1 genes in different rice tissues and under different stress have been analysised. In addition, over-expression vector and RNAi vector of OsDREBA5-1 and OsRAVl-1 genes have been constructed successfully. The main results are as following:
1. OsHspl7.0-OE transgenic rice plants were obtained via Agrobacterium-mediated transformation and confirmed at transcriptional level. The results of salt and drought stress treatment showed that OsHspl7.0-OE transgenic plants exhibited less damage symptoms and higher survival rate than WT. After stress treatments, the proline content increased more significantly in transgenic plants than in wild type plants. By contrast, the MDA content and ion leakage increased more significantly in wild type plants than in transgenic plants. These results indicated that the over-expression of OsHsp17.0 gene in rice seedlings increased tolerance to salt stress and drought stress.
2. After high and cold temperature treatments, both OsHsp17.0-OE transgenic and WT plants can keep on growing, and no significant phenotypic differences have been found. After stress treatments, the proline content increased more significantly in transgenic plants than in wild type plants. Oppositely, the MDA content increased more significantly in wild type plants than in transgenic plants.
3. After stress treatment with heavy metal ions (Cadmium), the growth of both OsHsp17.0-OE transgenic and WT seedlings were inhibited, but no significant phenotypic difference have been found.
4. The two AP2/EREBP family gene OsDREBA5-1 and OsRAVl-1 exhibited tissue-specific expressions under normal condition.OsDREBA5-1 gene is predominantly expressed in roots; while OsRAVl-1 has higher expression in leaves, suggesting that these two genes have different function emphasis.
5. OsDREBA5-1 and OsRAVl-1 are up-regulated by several stresses including high temperature, PEG stress, salt stress and ABA, however, differential expression profiles have been observed. It is interesting that, under streses, the induced expression of OsDREBA5-1 is much stronger than OsRAVl-1 in rice roots; by contrast, the induced expression of OsRAVl-1 is much stronger than OsDREBA5-1 in rice shoots. It is suggest that OsDREBA5-1 mainly function in the roots and OsRAVl-1 mainly function in the shoots.
6. Plant over-expression and RNAi inhibition vectors for the two AP2/EREBP gene OsDREBA5-1 and OsRAVl-1 were constructed. Transgenic rice plants were-obtained via Agrobacterium-mediated transformation and confirmed at transcriptional level.
1.采用农杆菌介导法将OsHsp17.0基因的过表达载体转入水稻,从转录水平证实获得了阳性植株。对转基因植株进行盐胁迫和干旱胁迫处理后发现,OsHsp17.0-OE转基因幼苗较野生型WT出现受害症状的时间延迟,存活率提高；胁迫处理后转基因植株中脯氨酸的含量增幅大于野生型植株的,而MDA含量和离子渗漏率则相反。这些结果说明过表达OsHsp17.0提高了转基因水稻苗期对盐胁迫和干旱胁迫的耐受能力。
2.对转基因植株进行高温和低温胁迫处理后发现,OsHsp17.0-OE转基因和野生型WT幼苗均可继续生长,但二者无明显表型差异；转基因植株中脯氨酸的含量增幅大于野生型植株的,而MDA含量则相反。
3.对转基因植株进行重金属离子(镉)胁迫处理后发现,OsHsp17.0-OE转基因和野生型WT幼苗生长均受到抑制,但二者无明显表型差异。
4.两个AP2/EREBP基因OsDREBA5-1和OsRAV1-1的表达具有组织特异性。相比之下,OsDREBA5-1基因更趋向于在根部表达,而OsRAV1-1基因在叶和叶鞘表达更为强烈,推测这两个基因的功能侧重点不同。
S.OsDREBA5.1和OsRAV1-1基因在高温、PEG、NaCl和ABA处理中表达量均上调,但其表达模式有所区别。在多种逆境处理下,OsDREBA5-1基因在根部被诱导的程度要大大高于OsRAV1-1基因被诱导的程度；而在地上部分的情形恰好相反,OsRAV1-1基因被诱导的程度大大高于OsDREBA5-1基因被诱导的程度。这与这两个基因的部位表达特点相一致。OsDREBA5-1基因可能侧重于在根部参与调控对逆境的响应；而OsRAV1-1基因则更主要地在地上部分参与调控对逆境的响应。
6.构建了OsDREBA5-1和OsRAV1-1基因的过表达和RNAi干扰载体,通过农杆菌介导法获得转基因植株,并从转录水平证实获得了阳性植株。为研究这两个基因在水稻中的功能奠定了基础。
Rice is an important crop that more than half of the world people live on. Abiotic stresses such as high temperature, drought, and high salinity affect rice growth and development resulting in serious influence on rice yield and quality performance. Therefore, it is very important to find key regulatory or functional genes responsive to abiotic stresses, to reveal the molecular mechanisms of stress tolerance and to improve rice resistance by genetic engineering techniques. Heat shock proteins (Hsps) are important proteins whose expression is increased when the cells are exposed to elevated temperatures or other stresses. Increased expression of these proteins can protect the organism against stress-induced damage. Small Heat shock proteins (sHsps) are more abundant in plant. The AP2/EREBP transcription factors are known to be unique in plants, and they are involved in growth, development and environmental stress responses. DREB and RAV are two major subfamilies of AP2/EREBP. To investigate the gene function of OsHsp17.0,OsDREBA5-1 and OsRAVl-1, OsHspl7.0-overexpression rice plants have been exposed to different abiotic stresses including salt stress, drought stress, high temperature and low temperature treatment and the stress-tolerance has been analysised. Using semiquantitative RT-PCR method and real-time RT-PCR, the expression profile of OsDREBA5-1 and OsRAVl-1 genes in different rice tissues and under different stress have been analysised. In addition, over-expression vector and RNAi vector of OsDREBA5-1 and OsRAVl-1 genes have been constructed successfully. The main results are as following:
1. OsHspl7.0-OE transgenic rice plants were obtained via Agrobacterium-mediated transformation and confirmed at transcriptional level. The results of salt and drought stress treatment showed that OsHspl7.0-OE transgenic plants exhibited less damage symptoms and higher survival rate than WT. After stress treatments, the proline content increased more significantly in transgenic plants than in wild type plants. By contrast, the MDA content and ion leakage increased more significantly in wild type plants than in transgenic plants. These results indicated that the over-expression of OsHsp17.0 gene in rice seedlings increased tolerance to salt stress and drought stress.
2. After high and cold temperature treatments, both OsHsp17.0-OE transgenic and WT plants can keep on growing, and no significant phenotypic differences have been found. After stress treatments, the proline content increased more significantly in transgenic plants than in wild type plants. Oppositely, the MDA content increased more significantly in wild type plants than in transgenic plants.
3. After stress treatment with heavy metal ions (Cadmium), the growth of both OsHsp17.0-OE transgenic and WT seedlings were inhibited, but no significant phenotypic difference have been found.
4. The two AP2/EREBP family gene OsDREBA5-1 and OsRAVl-1 exhibited tissue-specific expressions under normal condition.OsDREBA5-1 gene is predominantly expressed in roots; while OsRAVl-1 has higher expression in leaves, suggesting that these two genes have different function emphasis.
5. OsDREBA5-1 and OsRAVl-1 are up-regulated by several stresses including high temperature, PEG stress, salt stress and ABA, however, differential expression profiles have been observed. It is interesting that, under streses, the induced expression of OsDREBA5-1 is much stronger than OsRAVl-1 in rice roots; by contrast, the induced expression of OsRAVl-1 is much stronger than OsDREBA5-1 in rice shoots. It is suggest that OsDREBA5-1 mainly function in the roots and OsRAVl-1 mainly function in the shoots.
6. Plant over-expression and RNAi inhibition vectors for the two AP2/EREBP gene OsDREBA5-1 and OsRAVl-1 were constructed. Transgenic rice plants were-obtained via Agrobacterium-mediated transformation and confirmed at transcriptional level.
引文
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