植物应答环境信号的两个相关基因功能研究
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摘要
与动物不同,植物不能通过移动以规避环境变化带来的风险。然而,植物已进化出相应的应答机制,以应对外界环境的变化,即根据环境信号调控生长发育以实现对逆境环境的适应。因此,解析植物生长发育与环境信号响应的分子机制一直是植物分子生态学的重要研究领域之一
植物对外界环境信号的应答是通过细胞内外的系列蛋白介导完成的。其中,位于细胞膜上的受体蛋白激酶能感受细胞外的信号并启动精准的应答过程；锚蛋白(接头蛋白)在细胞内介导蛋白与蛋白的特异互作,通过锚定重复基序作“接头”把蛋白招募在一起,从而使植物能特异地对外界环境作出适宜的应答,并通过调控其生长发育实现对环境的适应。
本文以受体蛋白激酶FERONIA(FER)和锚蛋白ANK6的功能研究为切入点,力图揭示它们在植物响应外界生态环境中的作用以及其与植物生长发育的关系,以促进对植物适应环境的分子机制的认识。主要研究结果如下：
一、受体蛋白激酶FER在逆境激素信号转导中的作用
1、首次发现FER-GEF1/4/10-ROP11信号通路为ABA信号途径关键蛋白ABI2的正调节因子
ABA是植物响应逆境的关键激素分子。ABA受体蛋白(PYR/PYL/RCAR)作为负调节因子抑制PP2CA家族蛋白ABI2的磷酸酶活性已见报道。本研究首次鉴定出ABI2的正调节因子,即FER-GEF1/4/10-ROP11信号通路直接激活ABI2的磷酸酶活性,负调节植物对ABA的响应。该通路与ABA受体蛋白家族对PP2CA的负调节通路一起构成了PP2CA活性调节的“阴-阳”两面。研究还发现ABI2直接与FER互作,作为一种反馈抑制方式抑制FER活性。在此基础上,建立了一个以FER为中心的ABA响应调节网络。该网络对植物气孔开关调控机制的理解、逆境生长表型的解析都将起重要促进作用。
2、首次发现FER是乙烯底物合成关键酶SAM1/2的负调节因子
乙烯是植物逆境响应中的另一关键激素分子。SAM1/2是乙烯合成底物S-腺苷甲硫氨酸(SAM)的关键合成酶。本研究发现FER和SAM1、SAM2在细胞膜上相互作用,负调节SAM的合成,进而降低乙烯的合成,从而影响植物的逆境响应和生长发育。
3、提出以FER为节点的植物响应环境信号的激素交叉会话模型
根据环境条件的变化,FER通过影响ABA、乙烯、生长素等多种激素响应,实现激素交叉会话,调控多条细胞信号转导通路,使植物对盐、冷、热、渗透、病原菌等非生物胁迫和生物胁迫环境信号作出响应,并通过调控生长发育以适应环境。
4、初步揭示了两个泛素连接酶对FER信号通路的调节作用
发现两个同源性非常高的ARM-E3连接酶家族成员(FEP1、FEP2)与FER互作,并初步证明FEP1/2可能是FER的下游正响应成员负调节保卫细胞对ABA的响应。该研究结果对FER介导的环境信号响应机制的理解有促进作用。
5、初步揭示了FER调控种子大小的分子机制。
发现FER在球形胚时期的种皮细胞中高度极性表达,通过GEF1负调节种皮细胞伸长速度从而调控种子大小。
二、锚蛋白ANK6在植物双受精过程中的作用
锚蛋白是介导蛋白与蛋白互作的“接头蛋白”,可能在RLK等介导的环境信号响应中起作用。ANK6是一个表达量受ABA和盐胁迫抑制的锚蛋白。在探索ANK6逆境响应功能的过程中,发现无法获得ank6纯合子,其杂合体的果荚中出现和fer果荚中类似的空位现象,并且其死亡胚珠的外观和fer类似。经研究证明ANK6在线粒体中通过与转录起始因子SIG5互作控制植物的双受精过程。初步解析了ANK6通过RPS9在植物生殖发育、ABA及盐胁迫响应中的作用。遗憾的是有关ANK6作为接头蛋白在植物环境响应响应中的作用进展不大,还需作更艰辛地探讨。
Unlike animals, plants can not "walk away" from the harmful environment. However, they have evolved elaborate mechanisms to deal with the changes of outside environments. That is to say, plants can acquire their adaptation to stress environments by controlling growth and development on the basis of environmental signals. Therefore, it's one of the hottest research realms in plant molecular ecology to dissect molecular mechanisms between growth and development of plants and their responses to environmental signals.
Plants fulfill their responses to environmental signals through a series of intra-and extra-cellular proteins. Among them, the plasma membrane-located receptor-like protein kinases (RLKs) are able to sense extra-cellular signals, and switch on corresponding elaborate responses. Ankyrin repeat proteins (adaptor proteins) mediate protein-protein interactions in cells by recruiting proteins through ankyrin repeat motif "adaptor", thus enable plants to response specifically to the outer environmental changes, and acquire adaptation to environments by means of controlling growth development.
We chose the FERONIA (FER) receptor kinase and ANK6ankyrin repeat protein as the starting point, to analyze their functions in plant ecological environmental responses and relations to plant growth development. These will promote us to understand the molecular mechanisms that plants adapt to environments. The main conclusions are as follows:
First, functions of receptor like kinase FER in the stress hormones signal transduction
1. FER-GEF1/4/10-ROP11-ABI2pathway was identified as positive regulation signal of a core factor of ABA responses, ABI2for the first time.
ABA is an important stress hormone. There are reporters that the ABA receptor proteins can inhibit the PP2CA (ABI2) activity. We identified a pathway which is composed of FER-GEF1/4/10-ROP11positively regulating ABI2activity and inhibit the ABA response. Both this pathway and the PP2CA negatively regulate pathway formed the "Ying-Yang" faces of the PP2CA activity regulators. On the other hand, ABI2physically interacts with the FER kinase, suggesting a feedback inhibition of FER by dephosphorylation, and so characterized an ABA response circuit by using FER as a node. This signal circuit will help us better understanding mechanisms of the guard cell open/closure and stress response control.
2. FER was identified as negatively regulator of the ethylene substrate synthetase SAM1and SAM2for the first time
Ethylene is another stress response hormone. The S-adenosyl-L-methi- onine (SAM) synthetase (SAM1and SAM2) can synthesize SAM, the precursor of ethylene. We found that FER interacts with SAM1/2on the plasma membrane and negatively regulate SAM synthesis, thereby inhibits ethylene synthesis and stimulates plants stress and development response at last.
3. A model was proposed that FER functions as a key node of multiple hormone cross-talks for plant to response stress environmental signalings.
According to the environmental signals, FER can regulate many signaling pathways by affecting ABA, ethylene or auxin response, thus get plants to response to abiotic stresses (salt, hot, cold and osmotic stress) or biotic stress, and finally adapt to outer environment by controlling plant growth and development.
4. Function of two E3ligases in FER network was analyzed preliminarily.
We found that two E3ligase which have high homology can interact with FER and function as positive response factors of FER ABA control pathway. These data will help us understanding the FER network.
5. Molecular mechanism of FER in seed size control was analyzed
FER displayed high expression and polarity in integuments, and can negatively regulate the seed size through GEF1to inhibit the integument cells elongation.
Second, functional analysis of ANK6during fertilization
Ankyrin repeat protein is an "adaptor protein"and can mediate the protein-protein interaction and may have function in the environmental responses in the RLKs mediated pathways. We found that ANK6expression is inhibited by ABA and salt stress. We want to explore the function of ANK6during the environmental response. But the lack of homozygous mutant from a T-DNA insertional line indicated that loss-of-function of ANK6results in embryonic lethality which is very similar to the fer fertilization phenotype. ANK6was localized to the mitochondria where it interacts with SIG5, a transcription initiation factor previously found to be essential for fertility. Moreover, ANK6can interact with a small ribosome subunit protein RPS9in mitochondrial. Loss-of-function of RPS9results in male, female gametophytes development, ABA and salt stress response defect. Unfortunately, as the embryonic lethality limitations, we only analyzed the function of ANK6in fertilization but did not analyze its function in environmental responses. More efforts are required in environmental response study in future.
植物对外界环境信号的应答是通过细胞内外的系列蛋白介导完成的。其中,位于细胞膜上的受体蛋白激酶能感受细胞外的信号并启动精准的应答过程；锚蛋白(接头蛋白)在细胞内介导蛋白与蛋白的特异互作,通过锚定重复基序作“接头”把蛋白招募在一起,从而使植物能特异地对外界环境作出适宜的应答,并通过调控其生长发育实现对环境的适应。
本文以受体蛋白激酶FERONIA(FER)和锚蛋白ANK6的功能研究为切入点,力图揭示它们在植物响应外界生态环境中的作用以及其与植物生长发育的关系,以促进对植物适应环境的分子机制的认识。主要研究结果如下：
一、受体蛋白激酶FER在逆境激素信号转导中的作用
1、首次发现FER-GEF1/4/10-ROP11信号通路为ABA信号途径关键蛋白ABI2的正调节因子
ABA是植物响应逆境的关键激素分子。ABA受体蛋白(PYR/PYL/RCAR)作为负调节因子抑制PP2CA家族蛋白ABI2的磷酸酶活性已见报道。本研究首次鉴定出ABI2的正调节因子,即FER-GEF1/4/10-ROP11信号通路直接激活ABI2的磷酸酶活性,负调节植物对ABA的响应。该通路与ABA受体蛋白家族对PP2CA的负调节通路一起构成了PP2CA活性调节的“阴-阳”两面。研究还发现ABI2直接与FER互作,作为一种反馈抑制方式抑制FER活性。在此基础上,建立了一个以FER为中心的ABA响应调节网络。该网络对植物气孔开关调控机制的理解、逆境生长表型的解析都将起重要促进作用。
2、首次发现FER是乙烯底物合成关键酶SAM1/2的负调节因子
乙烯是植物逆境响应中的另一关键激素分子。SAM1/2是乙烯合成底物S-腺苷甲硫氨酸(SAM)的关键合成酶。本研究发现FER和SAM1、SAM2在细胞膜上相互作用,负调节SAM的合成,进而降低乙烯的合成,从而影响植物的逆境响应和生长发育。
3、提出以FER为节点的植物响应环境信号的激素交叉会话模型
根据环境条件的变化,FER通过影响ABA、乙烯、生长素等多种激素响应,实现激素交叉会话,调控多条细胞信号转导通路,使植物对盐、冷、热、渗透、病原菌等非生物胁迫和生物胁迫环境信号作出响应,并通过调控生长发育以适应环境。
4、初步揭示了两个泛素连接酶对FER信号通路的调节作用
发现两个同源性非常高的ARM-E3连接酶家族成员(FEP1、FEP2)与FER互作,并初步证明FEP1/2可能是FER的下游正响应成员负调节保卫细胞对ABA的响应。该研究结果对FER介导的环境信号响应机制的理解有促进作用。
5、初步揭示了FER调控种子大小的分子机制。
发现FER在球形胚时期的种皮细胞中高度极性表达,通过GEF1负调节种皮细胞伸长速度从而调控种子大小。
二、锚蛋白ANK6在植物双受精过程中的作用
锚蛋白是介导蛋白与蛋白互作的“接头蛋白”,可能在RLK等介导的环境信号响应中起作用。ANK6是一个表达量受ABA和盐胁迫抑制的锚蛋白。在探索ANK6逆境响应功能的过程中,发现无法获得ank6纯合子,其杂合体的果荚中出现和fer果荚中类似的空位现象,并且其死亡胚珠的外观和fer类似。经研究证明ANK6在线粒体中通过与转录起始因子SIG5互作控制植物的双受精过程。初步解析了ANK6通过RPS9在植物生殖发育、ABA及盐胁迫响应中的作用。遗憾的是有关ANK6作为接头蛋白在植物环境响应响应中的作用进展不大,还需作更艰辛地探讨。
Unlike animals, plants can not "walk away" from the harmful environment. However, they have evolved elaborate mechanisms to deal with the changes of outside environments. That is to say, plants can acquire their adaptation to stress environments by controlling growth and development on the basis of environmental signals. Therefore, it's one of the hottest research realms in plant molecular ecology to dissect molecular mechanisms between growth and development of plants and their responses to environmental signals.
Plants fulfill their responses to environmental signals through a series of intra-and extra-cellular proteins. Among them, the plasma membrane-located receptor-like protein kinases (RLKs) are able to sense extra-cellular signals, and switch on corresponding elaborate responses. Ankyrin repeat proteins (adaptor proteins) mediate protein-protein interactions in cells by recruiting proteins through ankyrin repeat motif "adaptor", thus enable plants to response specifically to the outer environmental changes, and acquire adaptation to environments by means of controlling growth development.
We chose the FERONIA (FER) receptor kinase and ANK6ankyrin repeat protein as the starting point, to analyze their functions in plant ecological environmental responses and relations to plant growth development. These will promote us to understand the molecular mechanisms that plants adapt to environments. The main conclusions are as follows:
First, functions of receptor like kinase FER in the stress hormones signal transduction
1. FER-GEF1/4/10-ROP11-ABI2pathway was identified as positive regulation signal of a core factor of ABA responses, ABI2for the first time.
ABA is an important stress hormone. There are reporters that the ABA receptor proteins can inhibit the PP2CA (ABI2) activity. We identified a pathway which is composed of FER-GEF1/4/10-ROP11positively regulating ABI2activity and inhibit the ABA response. Both this pathway and the PP2CA negatively regulate pathway formed the "Ying-Yang" faces of the PP2CA activity regulators. On the other hand, ABI2physically interacts with the FER kinase, suggesting a feedback inhibition of FER by dephosphorylation, and so characterized an ABA response circuit by using FER as a node. This signal circuit will help us better understanding mechanisms of the guard cell open/closure and stress response control.
2. FER was identified as negatively regulator of the ethylene substrate synthetase SAM1and SAM2for the first time
Ethylene is another stress response hormone. The S-adenosyl-L-methi- onine (SAM) synthetase (SAM1and SAM2) can synthesize SAM, the precursor of ethylene. We found that FER interacts with SAM1/2on the plasma membrane and negatively regulate SAM synthesis, thereby inhibits ethylene synthesis and stimulates plants stress and development response at last.
3. A model was proposed that FER functions as a key node of multiple hormone cross-talks for plant to response stress environmental signalings.
According to the environmental signals, FER can regulate many signaling pathways by affecting ABA, ethylene or auxin response, thus get plants to response to abiotic stresses (salt, hot, cold and osmotic stress) or biotic stress, and finally adapt to outer environment by controlling plant growth and development.
4. Function of two E3ligases in FER network was analyzed preliminarily.
We found that two E3ligase which have high homology can interact with FER and function as positive response factors of FER ABA control pathway. These data will help us understanding the FER network.
5. Molecular mechanism of FER in seed size control was analyzed
FER displayed high expression and polarity in integuments, and can negatively regulate the seed size through GEF1to inhibit the integument cells elongation.
Second, functional analysis of ANK6during fertilization
Ankyrin repeat protein is an "adaptor protein"and can mediate the protein-protein interaction and may have function in the environmental responses in the RLKs mediated pathways. We found that ANK6expression is inhibited by ABA and salt stress. We want to explore the function of ANK6during the environmental response. But the lack of homozygous mutant from a T-DNA insertional line indicated that loss-of-function of ANK6results in embryonic lethality which is very similar to the fer fertilization phenotype. ANK6was localized to the mitochondria where it interacts with SIG5, a transcription initiation factor previously found to be essential for fertility. Moreover, ANK6can interact with a small ribosome subunit protein RPS9in mitochondrial. Loss-of-function of RPS9results in male, female gametophytes development, ABA and salt stress response defect. Unfortunately, as the embryonic lethality limitations, we only analyzed the function of ANK6in fertilization but did not analyze its function in environmental responses. More efforts are required in environmental response study in future.
引文
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