大豆转录因子GmNAC2和GmNAC5功能验证
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摘要
NAC转录因子是植物中家族成员较多的转录因子之一,它的表达受发育时期和多种环境因素的诱导,并在植物生理、发育等过程中起到广泛的调控作用。大豆的NAC转录因子研究在近5年才开始发展,因而对其的调控机制还不是很明确。本文研究探讨了两个大豆转录因子GmNAC2和GmNAC5的功能,发现GmNAC2转录因子参与逆境调控,GmNAC5转录因子在大豆发育中参与调控,为大豆种子产量和品质的遗传改良提供理论依据。
GmNAC2属于NAC转录因子ATAF亚家族。ATAF亚家族包括许多参与生物胁迫或非生物胁迫应答过程的蛋白,例如ATAF1, ATAF2和OsNAC6蛋白等。组织表达分析显示,GmNAC2基因在正在发育的种子中(开花后40天)表达量最高,预示着GmNAC2转录因子可能参与种子的发育过程。GmNAC2基因在根中也有较高的表达量。同时逆境胁迫发现GmNAC2受JA、机械损伤、高盐、干旱胁迫和冷冻处理等非生物胁迫的诱导表达,这与GmNAC2在根中较高水平的组织表达有一定的关联。GmNAC2基因在100μMABA处理下表达量的变化不明显,这可能说明在GmNAC2在非生物逆境胁迫下的诱导表达并不依赖于ABA。
为分析GmNAC2转录因子在非生物逆境胁迫下的调控作用,我们将GmNAC2基因转化烟草和大豆毛状根。分析发现,GmNAC2基因过量表达烟草对干旱、高盐和冷害胁迫较敏感,表现出幼苗矮小,根长较短等现象,同时逆境下转基因烟草的MDA含量比野生型烟草高,说明转基因烟草的膜脂过氧化反应和活性氧毒害作用较强烈。GmNAC2基因过量表达的转基因烟草叶片进行NBT和DAB组织染色,也发现逆境下超氧阴离子自由基和过氧化氢增多。对转基因大豆毛状根进行干旱、高盐和冷害胁迫处理,发现过量表达GmNAC2基因导致大豆毛状根MDA含量升高,SOD酶活和APX酶活下降,而沉默GmNAC2基因导致大豆毛状根MDA含量降低,SOD酶活和APX酶活上升。以上这些现象表明GmNAC2转录因子使逆境条件下的转基因烟草和大豆毛状根维持活性氧产生与清除动态平衡的能力明显下降,造成较大的氧化伤害。
为进一步分析GmNAC2转录因子调控ROS平衡体系的机制,对正常条件下生长的GmNAC2基因过量表达转基因烟草叶片进行转录本分析,发现一些抗氧化相关的基因(涉及抗氧化酶类和非酶促的抗氧化物质)都发生下调现象,例如编码SOD酶的CSD2,编码硫氧还原蛋白的ACHT5,催化AsA产生的MIOX1和MIOX2等。结合上述GmNAC2转录因子使逆境条件下的转基因烟草和大豆毛状根氧化伤害增加这些现象,可以推断GmNAC2转录因子负调控与活性氧清除途径相关的一些基因,导致逆境条件下GmNAC2基因过量表达转基因植株活性氧增多,氧化伤害加剧。
GmNAC5属于NAM亚家族,目前已经报到的这个家族的成员主要维持和调节顶端分生组织的正常发育过程。组织表达分析表明,GmNAC5基因在正在发育的种子中(开花后40天)表达量最高,表明GmNAC5基因可能与种子的发育过程有关；GmNAC5基因在根中的表达量也较高。分析GmNAC5基因在非生物胁迫下的表达,发现GmNAC5受机械损伤、高盐和冷害诱导,预示GmNAC5转录因子可能也参与了胁迫代谢途径。同时GmNAC5基因在100μM ABA处理下表达量的变化不明显,这可能说明GmNAC5在非生物逆境胁迫下的诱导表达并不依赖于ABA。在研究中发现GmNAC5转基因烟草叶片形态发生改变,过量表达GmNAC5基因导致烟草叶片变狭长。同时通过农杆菌介导的大豆子叶节转化法,获得了GmNAC5基因过量表达转基因大豆,qPCR验证,GmNAC5基因在转基因大豆叶片中表达量显著提高(最高10倍左右的表达量),观察T1代转基因大豆植株株型时,发现有部分转基因大豆侧枝明显增多,株型发生了改变。
The NAC (NAM, ATAF1,2, CUC2) family proteins are one of the largest plant-specific families of transcription factors which comprise several sub-families, such as ATAF, NAP and NAM. NAC (NAM, ATAF1,2, and CUC2) proteins have regulatory activity, transcription regulation and resistance to stress in plant development. NAC genes may be induced by developmental events or multiple environmental factors. The research of NAC transcription factors of soybean (Glycine max (L.) Merr.) has developed only5years, so the regulation mechanism of most NAC proteins in soybean is not clear. Here we report functions of GmNAC2and GmNAC5to serve theory basis for genetic improvement of soybean quality.
GmNAC2(Glycine max NAC like gene2) is the first ATAF-like NAC identified transcription factor in soybean. The ATAF genes are involved in response to a variety of abiotic stresses, such as drought, cold and salinity. The soybean organ expression patterns of the gene were examined, and GmNAC2was highly expressed in the roots and immature embryos, especially strongly in immature embryo of40days after flowering. We found GmNAC2was induced by JA, mechanical wounding, high salinity, drought, and cold treatments, especially it was strongly affected by drought stress, but was not induced by ABA. The result suggests that up-regulation of GmNAC2by drought, salinity and cold may be not dependent on ABA.
To analyze regulatory effect of GmNAC2gene under abiotic stresses, the transgenic tobacco and hairy root of GmNAC2gene were used for further study. We found GmNAC2overexpression tobacco lines were hypersensitive to drought, high salinity, and cold stress. MDA content of transgenic tobacco was higher than CK under abiotic stresses while O2and H2O2(ROS) contents of transgenic tobacco were also higher than CK after stress treatments. GmNAC2overexpression hairy roots reduces SOD activities and APX activities under drought, salt and cold stress, and GmNAC2silencing hairy roots enhances SOD activities and APX activities under drought stress, salt stress and cold stress conditions. The results demonstrate the great potential of this gene to reduce the ROS-scavenging ability and stress tolerance of crops.
To further elucidate the mechanism of GmNAC2-mediated tolerance, we performed transcriptome analyses comparing GmNAC2overexpression tobacco with wild type tobacco leaves under normal conditions. We identified interesting down-regulated genes such as related to ROS-scavenging in GmNAC2overexpression tobacco. The down-regulated genes related to ROS-scavenging include genes coding thioredoxin, SOD enzyme, peroxiredoxin proteins. Collectively, GmNAC2may function as a negative regulator in drought, high salinity, and cold stress signaling pathways through modulation of ROS-scavenging genes expression.
GmNAC5(Glycine max NAC like gene5) is a member of NAM subfamily belonging to NAC transcription factor in soybean. To date, the documents about NAM transcription factors showed that NAM subfamily members often maintained SAM development. The soybean organ expression patterns of the gene were examined, and GmNAC5gene was highly expressed in the roots and immature embryos, especially strongly in immature embryo of40days after flowering. We found GmNAC5was induced by mechanical wounding, high salinity and cold treatments, but was not induced by ABA. The results suggest GmNAC5may have regulatory effect in abiotic stresses response. GmNAC5overexpression transgenic tobacco plants have GmNAC5gene expression tested by qPCR. Analysis the leaves growth of transgenic plants and CK we found leaves of GmNAC5overexpression tobacco were narrower than CK tobacco. We first got GmNAC5overexpression transgenic soybean plants with about10times GmNAC5overexpression in leaves. Analyze the plant type of transgenic soybean, we found some T1generation plants had more lateral shoots.
GmNAC2属于NAC转录因子ATAF亚家族。ATAF亚家族包括许多参与生物胁迫或非生物胁迫应答过程的蛋白,例如ATAF1, ATAF2和OsNAC6蛋白等。组织表达分析显示,GmNAC2基因在正在发育的种子中(开花后40天)表达量最高,预示着GmNAC2转录因子可能参与种子的发育过程。GmNAC2基因在根中也有较高的表达量。同时逆境胁迫发现GmNAC2受JA、机械损伤、高盐、干旱胁迫和冷冻处理等非生物胁迫的诱导表达,这与GmNAC2在根中较高水平的组织表达有一定的关联。GmNAC2基因在100μMABA处理下表达量的变化不明显,这可能说明在GmNAC2在非生物逆境胁迫下的诱导表达并不依赖于ABA。
为分析GmNAC2转录因子在非生物逆境胁迫下的调控作用,我们将GmNAC2基因转化烟草和大豆毛状根。分析发现,GmNAC2基因过量表达烟草对干旱、高盐和冷害胁迫较敏感,表现出幼苗矮小,根长较短等现象,同时逆境下转基因烟草的MDA含量比野生型烟草高,说明转基因烟草的膜脂过氧化反应和活性氧毒害作用较强烈。GmNAC2基因过量表达的转基因烟草叶片进行NBT和DAB组织染色,也发现逆境下超氧阴离子自由基和过氧化氢增多。对转基因大豆毛状根进行干旱、高盐和冷害胁迫处理,发现过量表达GmNAC2基因导致大豆毛状根MDA含量升高,SOD酶活和APX酶活下降,而沉默GmNAC2基因导致大豆毛状根MDA含量降低,SOD酶活和APX酶活上升。以上这些现象表明GmNAC2转录因子使逆境条件下的转基因烟草和大豆毛状根维持活性氧产生与清除动态平衡的能力明显下降,造成较大的氧化伤害。
为进一步分析GmNAC2转录因子调控ROS平衡体系的机制,对正常条件下生长的GmNAC2基因过量表达转基因烟草叶片进行转录本分析,发现一些抗氧化相关的基因(涉及抗氧化酶类和非酶促的抗氧化物质)都发生下调现象,例如编码SOD酶的CSD2,编码硫氧还原蛋白的ACHT5,催化AsA产生的MIOX1和MIOX2等。结合上述GmNAC2转录因子使逆境条件下的转基因烟草和大豆毛状根氧化伤害增加这些现象,可以推断GmNAC2转录因子负调控与活性氧清除途径相关的一些基因,导致逆境条件下GmNAC2基因过量表达转基因植株活性氧增多,氧化伤害加剧。
GmNAC5属于NAM亚家族,目前已经报到的这个家族的成员主要维持和调节顶端分生组织的正常发育过程。组织表达分析表明,GmNAC5基因在正在发育的种子中(开花后40天)表达量最高,表明GmNAC5基因可能与种子的发育过程有关；GmNAC5基因在根中的表达量也较高。分析GmNAC5基因在非生物胁迫下的表达,发现GmNAC5受机械损伤、高盐和冷害诱导,预示GmNAC5转录因子可能也参与了胁迫代谢途径。同时GmNAC5基因在100μM ABA处理下表达量的变化不明显,这可能说明GmNAC5在非生物逆境胁迫下的诱导表达并不依赖于ABA。在研究中发现GmNAC5转基因烟草叶片形态发生改变,过量表达GmNAC5基因导致烟草叶片变狭长。同时通过农杆菌介导的大豆子叶节转化法,获得了GmNAC5基因过量表达转基因大豆,qPCR验证,GmNAC5基因在转基因大豆叶片中表达量显著提高(最高10倍左右的表达量),观察T1代转基因大豆植株株型时,发现有部分转基因大豆侧枝明显增多,株型发生了改变。
The NAC (NAM, ATAF1,2, CUC2) family proteins are one of the largest plant-specific families of transcription factors which comprise several sub-families, such as ATAF, NAP and NAM. NAC (NAM, ATAF1,2, and CUC2) proteins have regulatory activity, transcription regulation and resistance to stress in plant development. NAC genes may be induced by developmental events or multiple environmental factors. The research of NAC transcription factors of soybean (Glycine max (L.) Merr.) has developed only5years, so the regulation mechanism of most NAC proteins in soybean is not clear. Here we report functions of GmNAC2and GmNAC5to serve theory basis for genetic improvement of soybean quality.
GmNAC2(Glycine max NAC like gene2) is the first ATAF-like NAC identified transcription factor in soybean. The ATAF genes are involved in response to a variety of abiotic stresses, such as drought, cold and salinity. The soybean organ expression patterns of the gene were examined, and GmNAC2was highly expressed in the roots and immature embryos, especially strongly in immature embryo of40days after flowering. We found GmNAC2was induced by JA, mechanical wounding, high salinity, drought, and cold treatments, especially it was strongly affected by drought stress, but was not induced by ABA. The result suggests that up-regulation of GmNAC2by drought, salinity and cold may be not dependent on ABA.
To analyze regulatory effect of GmNAC2gene under abiotic stresses, the transgenic tobacco and hairy root of GmNAC2gene were used for further study. We found GmNAC2overexpression tobacco lines were hypersensitive to drought, high salinity, and cold stress. MDA content of transgenic tobacco was higher than CK under abiotic stresses while O2and H2O2(ROS) contents of transgenic tobacco were also higher than CK after stress treatments. GmNAC2overexpression hairy roots reduces SOD activities and APX activities under drought, salt and cold stress, and GmNAC2silencing hairy roots enhances SOD activities and APX activities under drought stress, salt stress and cold stress conditions. The results demonstrate the great potential of this gene to reduce the ROS-scavenging ability and stress tolerance of crops.
To further elucidate the mechanism of GmNAC2-mediated tolerance, we performed transcriptome analyses comparing GmNAC2overexpression tobacco with wild type tobacco leaves under normal conditions. We identified interesting down-regulated genes such as related to ROS-scavenging in GmNAC2overexpression tobacco. The down-regulated genes related to ROS-scavenging include genes coding thioredoxin, SOD enzyme, peroxiredoxin proteins. Collectively, GmNAC2may function as a negative regulator in drought, high salinity, and cold stress signaling pathways through modulation of ROS-scavenging genes expression.
GmNAC5(Glycine max NAC like gene5) is a member of NAM subfamily belonging to NAC transcription factor in soybean. To date, the documents about NAM transcription factors showed that NAM subfamily members often maintained SAM development. The soybean organ expression patterns of the gene were examined, and GmNAC5gene was highly expressed in the roots and immature embryos, especially strongly in immature embryo of40days after flowering. We found GmNAC5was induced by mechanical wounding, high salinity and cold treatments, but was not induced by ABA. The results suggest GmNAC5may have regulatory effect in abiotic stresses response. GmNAC5overexpression transgenic tobacco plants have GmNAC5gene expression tested by qPCR. Analysis the leaves growth of transgenic plants and CK we found leaves of GmNAC5overexpression tobacco were narrower than CK tobacco. We first got GmNAC5overexpression transgenic soybean plants with about10times GmNAC5overexpression in leaves. Analyze the plant type of transgenic soybean, we found some T1generation plants had more lateral shoots.
引文
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