植物激素脱落酸通过诱导乙烯的生物合成抑制拟南芥主根生长
摘要
植物在整个生命过程中面临着各种各样的生物及非生物胁迫,其固着生长的特性决定了它们只能通过调节自身的生命活动来应对环境的变化。抗逆性就是植物在对环境逐步适应的过程中形成的。根器官作为植物体的重要组成部分,不仅承担着营养物质的吸收、输送和贮藏的功能,还能感受内源信号和外界环境刺激并作出反应,从而使植物可以更好的生存繁衍。尤其在土壤贫瘠或干旱等恶劣环境中,根的作用更为重要。
脱落酸(abscisic acid, ABA)作为一种重要的植物激素,参与调控植物生长发育的各个阶段,包括种子的休眠及萌发、幼苗生长、气孔运动及对环境胁迫的响应等。高浓度的ABA具有抑制植物主根生长的作用,但其分子机理目前尚不清晰。
据文献报道,ABA通过影响乙烯的信号转导抑制拟南芥主根的生长,但不影响乙烯的生物合成。然而,本论文的结果却证明：乙烯合成途径中ACC合酶(ACC synthases, ACSs)的竞争性抑制剂氨基乙氧基乙烯甘氨酸(aminoethoxyvinylglycine, AVG),可以消除ABA对根生长的抑制作用。另外,通过气相色谱分析的方法直接的证明ABA可以诱导乙烯合成。以上结果暗示ABA可能是通过诱导乙烯的合成抑制根的生长,并且ACC合酶可能是ABA作用的靶点。为了进一步验证这一猜想,接下来又检测了ACC合酶T-DNA插入多突变体CS16647(acs1-1acs2-1acs4-1acs5-2acs6-1acs7-1)、CS16649(acs2-1acs4-1acs5-2acs6-1acs7-1acs9-1)和CS16650(acs1-1acs2-1acs4-1acs5-2acs6-1acs7-1acs9-1)对ABA的响应,发现它们主根的生长都具有ABA不敏感的表型。
ACC合酶是乙烯合成途径中的关键酶和限速酶,一般情况下在植物体内含量很低且蛋白极不稳定。qRT-PCR检测表明转录水平可能不是ABA调控ACC合酶的主要原因,ACC合酶转录后水平的调控至关重要。据文献报道,一型和二型ACC合酶的羧基端含有预测的CDPK磷酸化位点,可能参与其转录后水平的调控。已知两个钙依赖的蛋白激酶CPK4和CPKll的单突变体及双突变体的根生长对ABA反应不敏感,并且本论文发现,在ABA的处理下单突变体及双突变体的乙烯释放量少于野生型。体外磷酸化实验证明,CPK4和CPK1l可以磷酸化ACC合酶,并且其磷酸化活性受ABA诱导。之后利用定点突变的方法,找到ACS6的四个可能的CDPK磷酸化位点。这些磷酸化位点的突变并不影响该酶的活性,而是影响了蛋白的稳定性。
为了进一步研究CDPK磷酸化位点的生理功能,我们构建了过表达ACS6WT, ACS6AAAA及ACS6DDDD的转基因植株。研究发现,体外模拟CDPK磷酸化状态的ACS6DDDD转基因植株释放出更多的乙烯,并且主根生长及种子萌发变绿都对ABA反应不敏感,这些表型可能是由于乙烯过量合成引起的。
综上所述,ABA通过蛋白激酶CPK4及CPK1l调控ACSs蛋白的稳定性,影响乙烯的合成,进而调控根的生长。这一研究,有助于进一步了解ABA调控根生长的分子机理,同时对研究ABA和乙烯两大植物激素之间的相互作用也具有重要意义。
Plants are faced with a variety of abiotic and biotic stresses in their entire life cycle. The sessile nature determines that they have to cope with the environmental changes only through adjusting their life activities. The plant resistance is formed in the process of gradually adaption to the environment. As an important organ, root not only bears the functions of absorption, transport and storage of nutrients, but also feels the endogenous signals and exogenous environmental stimuli, so that plants could grow and develop much better. Root morphogenesis directly affects the growth of the whole plant, therefore is closely related to yield. Especially in poor soil or stressful conditions, such as drought, the function of root is more important.
During the whole life of plants, phytohormones play crucial roles. They interact with each other and co-regulate the physiological processes of plants. Abscisic acid (ABA) as an important phytohormone, regulates the growth and development in plant. Over a long period, it is known that high concentrations of ABA inhibit root growth. However, the molecular mechanism by which ABA regulates root growth remains unclear.
In Arabidopsis, the previous studies showed that ABA inhibits root growth through ethylene signaling, but not ethylene biosynthesis. In this study, we found that ABA inhibits root growth by regulating ethylene biosynthesis in Arabidopsis. The ethylene biosynthesis inhibitor aminoethoxyvinyl glycine (AVG) reduced the inhibition of root growth by ABA, and multiple ACS (1-aminocyclopropane-1-carboxylate synthase) mutants were more insensitive to ABA in terms of root growth than was the wild type. In addition, it was shown that ABA induced the ethylene biosynthesis by the gas chromatography testing.
ACC synthases, the key enzymes of ethylene biosynthesis pathway, are less abundant and unstable. Type1and type2ACS isozymes carry the predictive CDPK phosphorylation sites, and previous study showed that phosphorylation at specific sites by CDPKs could stabilize the protein production. Here, we reported that two ABA-activated CDPK protein kinases, CPK4and CPK11, phosphorylated the C-terminus of ACS6and stabilized ACS6protein. The transgenic plants expressing ACS6mutant form that mimics the CDPK phosphorylation produce more ethylene than dose the wild type, moreover, were more insensitive to ABA in terms of root growth and seeds germination greening than was the wild type.
This study helps us to further understand the molecular mechanism by which ABA regulates root growth. It will also shed more light on the mechanisms of crosstalk between ABA and ethylene signaling.
脱落酸(abscisic acid, ABA)作为一种重要的植物激素,参与调控植物生长发育的各个阶段,包括种子的休眠及萌发、幼苗生长、气孔运动及对环境胁迫的响应等。高浓度的ABA具有抑制植物主根生长的作用,但其分子机理目前尚不清晰。
据文献报道,ABA通过影响乙烯的信号转导抑制拟南芥主根的生长,但不影响乙烯的生物合成。然而,本论文的结果却证明：乙烯合成途径中ACC合酶(ACC synthases, ACSs)的竞争性抑制剂氨基乙氧基乙烯甘氨酸(aminoethoxyvinylglycine, AVG),可以消除ABA对根生长的抑制作用。另外,通过气相色谱分析的方法直接的证明ABA可以诱导乙烯合成。以上结果暗示ABA可能是通过诱导乙烯的合成抑制根的生长,并且ACC合酶可能是ABA作用的靶点。为了进一步验证这一猜想,接下来又检测了ACC合酶T-DNA插入多突变体CS16647(acs1-1acs2-1acs4-1acs5-2acs6-1acs7-1)、CS16649(acs2-1acs4-1acs5-2acs6-1acs7-1acs9-1)和CS16650(acs1-1acs2-1acs4-1acs5-2acs6-1acs7-1acs9-1)对ABA的响应,发现它们主根的生长都具有ABA不敏感的表型。
ACC合酶是乙烯合成途径中的关键酶和限速酶,一般情况下在植物体内含量很低且蛋白极不稳定。qRT-PCR检测表明转录水平可能不是ABA调控ACC合酶的主要原因,ACC合酶转录后水平的调控至关重要。据文献报道,一型和二型ACC合酶的羧基端含有预测的CDPK磷酸化位点,可能参与其转录后水平的调控。已知两个钙依赖的蛋白激酶CPK4和CPKll的单突变体及双突变体的根生长对ABA反应不敏感,并且本论文发现,在ABA的处理下单突变体及双突变体的乙烯释放量少于野生型。体外磷酸化实验证明,CPK4和CPK1l可以磷酸化ACC合酶,并且其磷酸化活性受ABA诱导。之后利用定点突变的方法,找到ACS6的四个可能的CDPK磷酸化位点。这些磷酸化位点的突变并不影响该酶的活性,而是影响了蛋白的稳定性。
为了进一步研究CDPK磷酸化位点的生理功能,我们构建了过表达ACS6WT, ACS6AAAA及ACS6DDDD的转基因植株。研究发现,体外模拟CDPK磷酸化状态的ACS6DDDD转基因植株释放出更多的乙烯,并且主根生长及种子萌发变绿都对ABA反应不敏感,这些表型可能是由于乙烯过量合成引起的。
综上所述,ABA通过蛋白激酶CPK4及CPK1l调控ACSs蛋白的稳定性,影响乙烯的合成,进而调控根的生长。这一研究,有助于进一步了解ABA调控根生长的分子机理,同时对研究ABA和乙烯两大植物激素之间的相互作用也具有重要意义。
Plants are faced with a variety of abiotic and biotic stresses in their entire life cycle. The sessile nature determines that they have to cope with the environmental changes only through adjusting their life activities. The plant resistance is formed in the process of gradually adaption to the environment. As an important organ, root not only bears the functions of absorption, transport and storage of nutrients, but also feels the endogenous signals and exogenous environmental stimuli, so that plants could grow and develop much better. Root morphogenesis directly affects the growth of the whole plant, therefore is closely related to yield. Especially in poor soil or stressful conditions, such as drought, the function of root is more important.
During the whole life of plants, phytohormones play crucial roles. They interact with each other and co-regulate the physiological processes of plants. Abscisic acid (ABA) as an important phytohormone, regulates the growth and development in plant. Over a long period, it is known that high concentrations of ABA inhibit root growth. However, the molecular mechanism by which ABA regulates root growth remains unclear.
In Arabidopsis, the previous studies showed that ABA inhibits root growth through ethylene signaling, but not ethylene biosynthesis. In this study, we found that ABA inhibits root growth by regulating ethylene biosynthesis in Arabidopsis. The ethylene biosynthesis inhibitor aminoethoxyvinyl glycine (AVG) reduced the inhibition of root growth by ABA, and multiple ACS (1-aminocyclopropane-1-carboxylate synthase) mutants were more insensitive to ABA in terms of root growth than was the wild type. In addition, it was shown that ABA induced the ethylene biosynthesis by the gas chromatography testing.
ACC synthases, the key enzymes of ethylene biosynthesis pathway, are less abundant and unstable. Type1and type2ACS isozymes carry the predictive CDPK phosphorylation sites, and previous study showed that phosphorylation at specific sites by CDPKs could stabilize the protein production. Here, we reported that two ABA-activated CDPK protein kinases, CPK4and CPK11, phosphorylated the C-terminus of ACS6and stabilized ACS6protein. The transgenic plants expressing ACS6mutant form that mimics the CDPK phosphorylation produce more ethylene than dose the wild type, moreover, were more insensitive to ABA in terms of root growth and seeds germination greening than was the wild type.
This study helps us to further understand the molecular mechanism by which ABA regulates root growth. It will also shed more light on the mechanisms of crosstalk between ABA and ethylene signaling.
引文
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