水稻ONAC家族基因重叠表达特性及其在抗病抗逆中的功能研究
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摘要
植物在生长发育的过程中,经常会受到干旱、高盐、低温、病原菌等各种非生物和生物胁迫的影响。这些胁迫通常会造成植物细胞的损伤,进而严重影响植物的生长发育和作物产量。在长期的进化过程中,植物形成了一系列生理、生化、代谢及防御机制,以适应和抵御各种生物和非生物逆境。在众多的适应和抗逆机制中,基因表达的转录调控在植物适应环境和抵御逆境胁迫中起重要作用。NAC (NAM/ATAF/CUC)家族基因是植物特有的一类转录因子,存在于多种植物中,其共同特点是在N端含有一段高度保守、约由150个氨基酸组成的NAC结构域,而C端为高度变异的转录调控区。研究表明,NAC转录因子不仅参与植物生长发育的调控、而且在植物抗病抗逆反应中具有重要的调控作用。本研究系统分析了ONAC家族基因对生物与非生物胁迫的响应、并分离克隆21个抗病抗逆相关的(?)NAC基因。在此基础上,研究了这21个ONAC基因在抗病信号分子处理、水稻-稻瘟菌和水稻-白叶枯病菌互作中的表达模式,以及初步分析了部分基因的生化特性和在抗病抗逆中的功能,旨在发现潜在的抗病抗逆胁迫相关的基因,为基因工程提高水稻抗病抗逆性能提供理论基础。本研究的主要内容及结果如下：
本研究利用水稻基因芯片公共数据库[Rice Oligonucleotide Array Database (ROAD)(http://www.ricearray.org/)],对多种生物[稻瘟病(Magnaporthe grisea)、寄生植物独脚金(Striga hermonthica)、水稻条纹病毒(Rice stripe virus, RSV)、水稻白叶枯病((?)(anthomonas oryzae pv.oryzae, Xoo)、水稻细菌性条斑病菌(Xanthomonas oryzae pv. oryzicola, Xoc)]与非生物(高盐、干旱、低温)胁迫下ONAC家族基因的响应进行分析,并选择30个ONAC基因对部分胁迫因子下的响应进行验证,结果发现该家族基因有63个ONAC基因对多种胁迫因子有重叠表达的特性,在这些重叠表达的基因中,多数基因在高盐、干旱、低温、稻瘟病、寄生植物独脚金、水稻条纹病毒诱导上调表达。对于非生物胁迫,38个ONAC基因重叠表达于任意两种胁迫条件,且选择的30个ONAC基因中有16个受脱落酸(Abscisic acid、ABA)诱导表达；而对于生物胁迫,有65个基因重叠表达于任意两种生物肋迫因子。系统进化树分析表明这些重叠表达的基因分布于ONAC家族的10个亚组[OsNAC7(SND)、ONAC11(NAC1)、 SNACⅠ、SNACⅡ (NAM/CUC3)、SNACⅢ(TIP)、SNACⅣ (ANAC34)、ONAC079(ONAC6、ONAC012(ONAC7)、ONAC135(0NAC3)和ONAC100(ONAC2)],同源搜索发现在该家族有16组基因高度同源。ONAC基因对多种胁迫因子的重叠表达特性及其基因间的高度同源性,预示着ONAC基因的功能多效性和相似性。
选择克隆了21个水稻抗病抗逆相关候选ONAC基因,研究了这些ONAC基因在抗病信号分子处理、水稻-稻瘟菌以及水稻-白叶枯病菌互作中的表达模式。结果表明分离获得的21个抗病抗逆相关ONAC基因中,除了ONAC058和(ONAC066,多数ONAC基因的表达能被水杨酸(Salicylic acid, SA)、茉莉酸(Jasmonic acid, JA)、 ACC (Ethylene, ET)多个抗病信号分子所诱导,但表达模式在不同抗病信号分子处理后不尽相同,相对于SA,多数ONAC基因对ACC (ET)或JA的响应程度较强。其中,12个ONAC基因(OsNAC8、ONAC016、ONAC022、ONAC023、ONAC049、 ONAC054、ONAC061、ONAC063、ONAC075、ONAC095、ONAC122和(ONAC131)的表达可以被ACC (ET)强烈诱导,6个ONAC基因(SNAC1、OsNAC4、OsNAC5、 OsNAC6、ONAC103和ONAC121)表达受JA强烈诱导,这些结果表明这18个抗病抗逆相关ONAC基因是通过JA/ET信号途径或通过JA/ET与SA信号途径参与水稻的抗病反应。在水稻-稻瘟菌、水稻-白叶枯病菌互作模式下,11个ONAC (ONAC022、ONAC023、ONAC049、ONAC054、ONAC061、ONAC075、ONAC095、 ONAC103、ONAC121、ONAC122和ONAC131)基因在水稻-稻瘟病菌、水稻-白叶枯病菌的非亲和性互作中特异性地诱导表达,因而可能与水稻抗稻瘟病、水稻白叶枯病反应的激活有关。此外,3个ONAC基因(OsNAC6、OsNAC8、ONAC063)在水稻-稻瘟病菌的非亲和性互作中特异性地诱导表达,而在水稻-白叶枯病菌的非亲和性互作中表达下调,这3个基因可能与抗水稻稻瘟病激活有关,且与抗水稻白叶枯病的抑制有关。4个ONAC基因(OsNAC3、OsNAC4、ONAC016和ONAC058)在水稻-白叶枯病菌非亲和性互作中上调表达,可能参与水稻对白叶枯抗病的激活有关。
选择研究5个ONAC基因(ONAC023、ONAC063、ONAC095、ONAc122、 ONAC131)的亚细胞定位和转录活性,发现这5个基因编码的蛋白都能定位于细胞核,且都具有转录激活活性。
采用病毒诱导的基因沉默技术(VIGS)初步对一些分离的ONAC基因的抗病功能进行初步的研究。结果发现当5个ONAC基因(ONAC023、ONAC095、ONAC122、 ONAC131(?)口(OsNAC6)分别沉默后,水稻抗稻瘟病害表现出感病性的增加,且一些抗病相关的基因(Oseds1、OsPAD4、OsLOX、OsPR3、OsPR1a、OsWRKY45、OsJamyb和(?)OsNH1/NPR1)也不同程度的表达下调,表明这5个ONAC基因通过直接或间接调控这些抗病相关的基因参与水稻对稻瘟病抗性。另外,ONAC023和ONAC095两个基因沉默后,对白叶枯病的抗病性下降,表明这两个基因可能在白叶枯抗病中具有正向调控的作用。
为了深入的探讨ONAC基因在抗病抗逆反应中的功能,构建了多数克隆的ONAC基因的超表达载体和SRDX载体,并导入水稻秀水11品种中。目前已经获得一些基因的超表达转基因株系或SRDX转基因株系,正在通过分子生物学方法对转基因株系进行鉴定,其中ONAC075-oe、OsNAC6-oe转基因株系的T0代种子已获得,并对其在抗病抗逆中的功能进行初步分析。结果表明过量表达OsNAC6和ONAC075的水稻转基因株系,对稻瘟病抗性和干旱胁迫的耐受性增加,而这2个基因参与植物抗病、抗逆的机制还需进一步的研究。
Plants are subjected to various types of abiotic (e.g. high-salinity, drought, and cold) and biotic (e.g. pathogen infection, insect fedding, and parasitic plants) stresses in their growth and development environments. To survive under such environment, plants have developed a series of mechanisms at physiological, biochemical, and molecular levels to adapt and defend themselves against these stresses. During these processes, different types of transcription factors play critical roles in plant defense response to biotic and abiotic stresses through directly or indirectly regulation of stress-related genes. NAC (NAM/ATAF/CUC) transcription factor, as one of largest and plant-specific gene families exsists in many different plants. The common characteristics of the NAC protein are the presence of a conserved NAC domain, composing of about150amino acids in N-terminals, and a highly variable transcriptional activation region in C-terminals. Extensive studies have revealed that NAC transcription factors not only play important roles in plant growth and development, but also have functions in regulation of responses to biotic and abiotic stresses. The study systematically investigated the expression characterization of ONAC genes under the different biotic and abiotic stresses. Based on the overlapping expression characteization, we isolated21ONAC genes from rice and investigated the expression patterns of these21ON AC genes after pathogens infection, and exogenous hormones salicylic acid (SA), methyl jasmonate (MeJA), ACC (ethylene,ET). The molecular characterization and the function of some ONAC genes were also investigated in this study. All of these studies will provide potential utility in rice genetic improvement of stress resistacne in rice.The major results of ONAC genes in this study were showed in the following aspects:
Microarray data analyses revealed that about63ONAC genes were overlapping expression during in various abiotic (salinity, drought, and Cold) and biotic stress (Magnaporthe grisea, Striga hermonthica, Xoo, Xoc, and RSV) conditions. Most of ONAC genes were up-regulated expression response to salt, dry, cold, Magnaporthe grisea, Striga hermonthica,and RSV. For the abotic stresses.38genes were overlapping expression in any two stress conditions.16among30representative ONAC genes were up-regulated expression response to Abscisic acid (ABA). For the biotic stresses,65ONAC genes were overlapping expression in any two biotic stresses. The phylogenetic tree suggested that these ONAC genes were classified into10subgroups [OsNAC7(SND), ONAC11(NAC1), SNACⅠ, SNACⅡ (NAM/CUC3), SNACⅢ (TIP), SNACⅣ (ANAC34), ONAC079(ONAC6), ONAC012(ONAC7), ONAC135(ONAC3). ONAC100(ONAC2)]. Homology sequence search found that there were many highly ortholgous genes in ONAC genes family. The overlapping expression of ONAC genes indicated that ONAC genes might play important roles in the cross-talk of different abiotic and biotic stresses and might have the multiple biological functions for the stress conditions.
We isolated21ONAC genes from rice and investigated the expression patterns of these21ONAC genes after pathogens infection, and exogenous hormones SA, MeJA, ACC. For pathogens infection, two interaction systems of rice-Magnaporthe grisea and rice-Xanthomonas oryzae pv.oryzae (Xoo) were used. The results showed that most of them were induced by the SA, JA and ACC (ET) except ONAC066and ONAC058. Among these,12ONAC genes(OsNAC8,ONAC016,ONAC022,ONAC023,ONAC049, ONAC054,ONAC061,ONAC063,ONAC075,ONAC095,ONAC122,and ONAC131) were strongly induced by ACC(ET),6ONAC genes (SNAC1,OsNAC4,OsNAC5,OsNAC6, ONAC103, and ONAC121) were strongly induced by JA. The expression patterns of21isolated ONAC genes in incompatible and compatible combinations between rice and Magnaporthe grisea, rice and Xoo were analyzed. The results showed that11ONAC genes (ONAC022, ONAC023, ONAC049, ONAC054, ONAC061, ONAC075, ONAC095, ONAC103, ONAC121, ONAC122, and ONAC131) were more strongly induced in the incompatible interaction than compatible interaction in these two combinations. According to the expression patterns, these11ONAC genes may be involved in resistance to fungal blast disease and bacterial blight of rice. In addition,3ONAC genes (OsNAC6, OsNAC8, and ONAC063) may be involved in resistance to fungal blast disease and susceptivity to Bacterial blight of rice, and4ONAC genes (OsNAC3, OsNAC4, ONAC016, and ONAC058) may be involved in resistance to Bacterial blight of rice.
Transient expression in tobacco epidermal cells indicated that ONAC-GFP fusion proteins including ONAC023, ONAC063, ONAC095, ONAC122, and ONAC131were localized in the nucleus. Transactivation analysis showed that ONAC023, ONAC063. ONAC095, ONAC122, or ONAC131played as a role in the activator of transcription.
We investigated the function of some ONAC genes in rice through virus-induced gene silencing (VIGS). The results showed that some pathogen relative genes (Oseds1, OsPAD4, OsLOX, OsPR3, OsPR1a, OsWRKY45, OsJamyb, and OsNHl/NPR1) were downregulation when ONAC023,ONAC095, ONAC122, ONAC131, or OsNAC6was silenced respectively, and finally the rice became more susceptive to rice blast.These results indicated that ONAC023, ONAC095, ONAC122, ONAC131, and OsNAC6were related to the resistance to rice blast disease. In addition, the rice seedling became more susceptive to rice bacterial blight when ONAC023or ONAC095was silenced respectively, which indicated that ONAC023and ONAC095play the positive roles in resistance to rice bacterial blitht.
To study the function of ONAC genes in disease resistance and stress tolerance, we constructed overexpression vectors (ONAC-OE) and chimeric repressor vectors (ONAC-SRDX) of isolated ONAC genes, and introduced them into wild-type rice XiuShui-11. In the present study, we have obtained some transgenic lines of ONAC genes and To seeds of ONAC075-oe, OsNAC6-oe transgenic lines. Furthermore, the function analysis of ONAC075-oe, OsNAC6-oe transgenic lines showed that ONAC075-oe and OsNAC6-oe can improve the blast disease resistance and drought stress tolerance.
本研究利用水稻基因芯片公共数据库[Rice Oligonucleotide Array Database (ROAD)(http://www.ricearray.org/)],对多种生物[稻瘟病(Magnaporthe grisea)、寄生植物独脚金(Striga hermonthica)、水稻条纹病毒(Rice stripe virus, RSV)、水稻白叶枯病((?)(anthomonas oryzae pv.oryzae, Xoo)、水稻细菌性条斑病菌(Xanthomonas oryzae pv. oryzicola, Xoc)]与非生物(高盐、干旱、低温)胁迫下ONAC家族基因的响应进行分析,并选择30个ONAC基因对部分胁迫因子下的响应进行验证,结果发现该家族基因有63个ONAC基因对多种胁迫因子有重叠表达的特性,在这些重叠表达的基因中,多数基因在高盐、干旱、低温、稻瘟病、寄生植物独脚金、水稻条纹病毒诱导上调表达。对于非生物胁迫,38个ONAC基因重叠表达于任意两种胁迫条件,且选择的30个ONAC基因中有16个受脱落酸(Abscisic acid、ABA)诱导表达；而对于生物胁迫,有65个基因重叠表达于任意两种生物肋迫因子。系统进化树分析表明这些重叠表达的基因分布于ONAC家族的10个亚组[OsNAC7(SND)、ONAC11(NAC1)、 SNACⅠ、SNACⅡ (NAM/CUC3)、SNACⅢ(TIP)、SNACⅣ (ANAC34)、ONAC079(ONAC6、ONAC012(ONAC7)、ONAC135(0NAC3)和ONAC100(ONAC2)],同源搜索发现在该家族有16组基因高度同源。ONAC基因对多种胁迫因子的重叠表达特性及其基因间的高度同源性,预示着ONAC基因的功能多效性和相似性。
选择克隆了21个水稻抗病抗逆相关候选ONAC基因,研究了这些ONAC基因在抗病信号分子处理、水稻-稻瘟菌以及水稻-白叶枯病菌互作中的表达模式。结果表明分离获得的21个抗病抗逆相关ONAC基因中,除了ONAC058和(ONAC066,多数ONAC基因的表达能被水杨酸(Salicylic acid, SA)、茉莉酸(Jasmonic acid, JA)、 ACC (Ethylene, ET)多个抗病信号分子所诱导,但表达模式在不同抗病信号分子处理后不尽相同,相对于SA,多数ONAC基因对ACC (ET)或JA的响应程度较强。其中,12个ONAC基因(OsNAC8、ONAC016、ONAC022、ONAC023、ONAC049、 ONAC054、ONAC061、ONAC063、ONAC075、ONAC095、ONAC122和(ONAC131)的表达可以被ACC (ET)强烈诱导,6个ONAC基因(SNAC1、OsNAC4、OsNAC5、 OsNAC6、ONAC103和ONAC121)表达受JA强烈诱导,这些结果表明这18个抗病抗逆相关ONAC基因是通过JA/ET信号途径或通过JA/ET与SA信号途径参与水稻的抗病反应。在水稻-稻瘟菌、水稻-白叶枯病菌互作模式下,11个ONAC (ONAC022、ONAC023、ONAC049、ONAC054、ONAC061、ONAC075、ONAC095、 ONAC103、ONAC121、ONAC122和ONAC131)基因在水稻-稻瘟病菌、水稻-白叶枯病菌的非亲和性互作中特异性地诱导表达,因而可能与水稻抗稻瘟病、水稻白叶枯病反应的激活有关。此外,3个ONAC基因(OsNAC6、OsNAC8、ONAC063)在水稻-稻瘟病菌的非亲和性互作中特异性地诱导表达,而在水稻-白叶枯病菌的非亲和性互作中表达下调,这3个基因可能与抗水稻稻瘟病激活有关,且与抗水稻白叶枯病的抑制有关。4个ONAC基因(OsNAC3、OsNAC4、ONAC016和ONAC058)在水稻-白叶枯病菌非亲和性互作中上调表达,可能参与水稻对白叶枯抗病的激活有关。
选择研究5个ONAC基因(ONAC023、ONAC063、ONAC095、ONAc122、 ONAC131)的亚细胞定位和转录活性,发现这5个基因编码的蛋白都能定位于细胞核,且都具有转录激活活性。
采用病毒诱导的基因沉默技术(VIGS)初步对一些分离的ONAC基因的抗病功能进行初步的研究。结果发现当5个ONAC基因(ONAC023、ONAC095、ONAC122、 ONAC131(?)口(OsNAC6)分别沉默后,水稻抗稻瘟病害表现出感病性的增加,且一些抗病相关的基因(Oseds1、OsPAD4、OsLOX、OsPR3、OsPR1a、OsWRKY45、OsJamyb和(?)OsNH1/NPR1)也不同程度的表达下调,表明这5个ONAC基因通过直接或间接调控这些抗病相关的基因参与水稻对稻瘟病抗性。另外,ONAC023和ONAC095两个基因沉默后,对白叶枯病的抗病性下降,表明这两个基因可能在白叶枯抗病中具有正向调控的作用。
为了深入的探讨ONAC基因在抗病抗逆反应中的功能,构建了多数克隆的ONAC基因的超表达载体和SRDX载体,并导入水稻秀水11品种中。目前已经获得一些基因的超表达转基因株系或SRDX转基因株系,正在通过分子生物学方法对转基因株系进行鉴定,其中ONAC075-oe、OsNAC6-oe转基因株系的T0代种子已获得,并对其在抗病抗逆中的功能进行初步分析。结果表明过量表达OsNAC6和ONAC075的水稻转基因株系,对稻瘟病抗性和干旱胁迫的耐受性增加,而这2个基因参与植物抗病、抗逆的机制还需进一步的研究。
Plants are subjected to various types of abiotic (e.g. high-salinity, drought, and cold) and biotic (e.g. pathogen infection, insect fedding, and parasitic plants) stresses in their growth and development environments. To survive under such environment, plants have developed a series of mechanisms at physiological, biochemical, and molecular levels to adapt and defend themselves against these stresses. During these processes, different types of transcription factors play critical roles in plant defense response to biotic and abiotic stresses through directly or indirectly regulation of stress-related genes. NAC (NAM/ATAF/CUC) transcription factor, as one of largest and plant-specific gene families exsists in many different plants. The common characteristics of the NAC protein are the presence of a conserved NAC domain, composing of about150amino acids in N-terminals, and a highly variable transcriptional activation region in C-terminals. Extensive studies have revealed that NAC transcription factors not only play important roles in plant growth and development, but also have functions in regulation of responses to biotic and abiotic stresses. The study systematically investigated the expression characterization of ONAC genes under the different biotic and abiotic stresses. Based on the overlapping expression characteization, we isolated21ONAC genes from rice and investigated the expression patterns of these21ON AC genes after pathogens infection, and exogenous hormones salicylic acid (SA), methyl jasmonate (MeJA), ACC (ethylene,ET). The molecular characterization and the function of some ONAC genes were also investigated in this study. All of these studies will provide potential utility in rice genetic improvement of stress resistacne in rice.The major results of ONAC genes in this study were showed in the following aspects:
Microarray data analyses revealed that about63ONAC genes were overlapping expression during in various abiotic (salinity, drought, and Cold) and biotic stress (Magnaporthe grisea, Striga hermonthica, Xoo, Xoc, and RSV) conditions. Most of ONAC genes were up-regulated expression response to salt, dry, cold, Magnaporthe grisea, Striga hermonthica,and RSV. For the abotic stresses.38genes were overlapping expression in any two stress conditions.16among30representative ONAC genes were up-regulated expression response to Abscisic acid (ABA). For the biotic stresses,65ONAC genes were overlapping expression in any two biotic stresses. The phylogenetic tree suggested that these ONAC genes were classified into10subgroups [OsNAC7(SND), ONAC11(NAC1), SNACⅠ, SNACⅡ (NAM/CUC3), SNACⅢ (TIP), SNACⅣ (ANAC34), ONAC079(ONAC6), ONAC012(ONAC7), ONAC135(ONAC3). ONAC100(ONAC2)]. Homology sequence search found that there were many highly ortholgous genes in ONAC genes family. The overlapping expression of ONAC genes indicated that ONAC genes might play important roles in the cross-talk of different abiotic and biotic stresses and might have the multiple biological functions for the stress conditions.
We isolated21ONAC genes from rice and investigated the expression patterns of these21ONAC genes after pathogens infection, and exogenous hormones SA, MeJA, ACC. For pathogens infection, two interaction systems of rice-Magnaporthe grisea and rice-Xanthomonas oryzae pv.oryzae (Xoo) were used. The results showed that most of them were induced by the SA, JA and ACC (ET) except ONAC066and ONAC058. Among these,12ONAC genes(OsNAC8,ONAC016,ONAC022,ONAC023,ONAC049, ONAC054,ONAC061,ONAC063,ONAC075,ONAC095,ONAC122,and ONAC131) were strongly induced by ACC(ET),6ONAC genes (SNAC1,OsNAC4,OsNAC5,OsNAC6, ONAC103, and ONAC121) were strongly induced by JA. The expression patterns of21isolated ONAC genes in incompatible and compatible combinations between rice and Magnaporthe grisea, rice and Xoo were analyzed. The results showed that11ONAC genes (ONAC022, ONAC023, ONAC049, ONAC054, ONAC061, ONAC075, ONAC095, ONAC103, ONAC121, ONAC122, and ONAC131) were more strongly induced in the incompatible interaction than compatible interaction in these two combinations. According to the expression patterns, these11ONAC genes may be involved in resistance to fungal blast disease and bacterial blight of rice. In addition,3ONAC genes (OsNAC6, OsNAC8, and ONAC063) may be involved in resistance to fungal blast disease and susceptivity to Bacterial blight of rice, and4ONAC genes (OsNAC3, OsNAC4, ONAC016, and ONAC058) may be involved in resistance to Bacterial blight of rice.
Transient expression in tobacco epidermal cells indicated that ONAC-GFP fusion proteins including ONAC023, ONAC063, ONAC095, ONAC122, and ONAC131were localized in the nucleus. Transactivation analysis showed that ONAC023, ONAC063. ONAC095, ONAC122, or ONAC131played as a role in the activator of transcription.
We investigated the function of some ONAC genes in rice through virus-induced gene silencing (VIGS). The results showed that some pathogen relative genes (Oseds1, OsPAD4, OsLOX, OsPR3, OsPR1a, OsWRKY45, OsJamyb, and OsNHl/NPR1) were downregulation when ONAC023,ONAC095, ONAC122, ONAC131, or OsNAC6was silenced respectively, and finally the rice became more susceptive to rice blast.These results indicated that ONAC023, ONAC095, ONAC122, ONAC131, and OsNAC6were related to the resistance to rice blast disease. In addition, the rice seedling became more susceptive to rice bacterial blight when ONAC023or ONAC095was silenced respectively, which indicated that ONAC023and ONAC095play the positive roles in resistance to rice bacterial blitht.
To study the function of ONAC genes in disease resistance and stress tolerance, we constructed overexpression vectors (ONAC-OE) and chimeric repressor vectors (ONAC-SRDX) of isolated ONAC genes, and introduced them into wild-type rice XiuShui-11. In the present study, we have obtained some transgenic lines of ONAC genes and To seeds of ONAC075-oe, OsNAC6-oe transgenic lines. Furthermore, the function analysis of ONAC075-oe, OsNAC6-oe transgenic lines showed that ONAC075-oe and OsNAC6-oe can improve the blast disease resistance and drought stress tolerance.
引文
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