高级植物生理学,信号转导

高级植物生理学,信号转导 高级植物生理学，植物细胞信号转导 植物在整个生长过程中，受到来自外部内部的各种生物的或非生物的刺激。如温度、光、重力、植物激素、病原生物，这些刺激都能影响植物的生长和发育过程。 在长期的进化过程中，植物体本身形成了许多应答(response)这些刺激的机制，其中信号转导(signal transduction)就是应答这些刺激最为重要的环节。细胞通过感受这些物理的、化学的或生物的信息，并作出适当的生理反应以维持其生命活动的进行。 例如植物的向光性能促使植物向光线充足的方向生长，在这个过程中，首先植物体要能感受到光线，然后把相关的信息传递到有关的靶细胞，并诱发胞内信号转化传递，调节基因的表达或改变酶的活性，从而使细胞作出反应。这种信息的胞间传递和胞内转导过程称为植物体内的信号传导。 植物体的信息系统可概括为两大类: 一类是遗传基因信息系统。在生物进化过程中形成的，以核酸蛋白质为组成的生物大分子信息系统。 高等植物细胞全能性的发现，证实了细胞中含有发育成为完整植株的全套的遗传信息。植物生长发育就是植物基因(遗传信息)受内外环境的影响，在时间和空间顺序表达的结果。 另一类是环境刺激—细胞反应偶联信息系统。植物感受到环境信号，并将其转变为植物体内的信号，从而调节植物的生长发育过程。 生长发育是基因在一定时间、空间上顺序表达的过程，而基因表达除受遗传信息支配外，还受环境的调控。 植物在整个生长发育过程中，受到各种内外因素的影响，植物体要正确地辨别接受各种信息并作出相应的反应，以确保正常生长和发育。 植物细胞受到的刺激或信号: 环境的外源信号: 如光照、温度、水分、重力、风、雨、气体、触摸、机械obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 伤害、病原因子等; 细胞的内源信号: 如激素、化学调节因子、电信号(生物电位)等。 这些信号作用于植物体，先到达细胞间隙—胞间信号，再到达细胞表面与细胞受体结合，通过跨膜信号转换，转变为胞内信号，将信息转导到胞内的特定效应部位起作用，而产生细胞反应，调节植物体的生长发育。 概念: 植物细胞信号转导(signal transduction)是指细胞感受各种刺激信号(包括各种内外刺激信号)与其引起特定生理效应的一系列分子反应机制。胞间信号分子又称第一信使，而胞内信号分子又称第二信使(是由胞外刺激信号引起改变的具有生理调节活性的细胞内因子)。 总之信号转导:就是植物感受环境因子(信号)后，通过不同途径、环节，一步步传递、转化，调节细胞中每个反应过程，产生相应的分子水平、生理生化代谢变化、及形态反应。 发展历史: , 20世纪60年代，提出环核苷酸(cAMP)为第二信使学说(胞间到胞内) , 70年代，Ca2+受体蛋白——钙调素(CaM)的发现，Ca2+作为第二信使 学说 , 80年代，质膜肌醇磷脂代谢途径产生的另外两个胞内信使——IP3和DG 的发现 , 同时，G蛋白的发现，依赖胞内信使的蛋白质可逆磷酸化的研究 信号传导的途径: , 胞间信号的感知和跨膜转换 , 胞内信号传导 , 蛋白质可逆磷酸化 , 细胞的生理反应 一、胞间信号感知和跨膜转换 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. (一)胞外信号的传递 1. 化学信号(chemical signals):是细胞感受刺激后合成并传递到作用部位引起生理反应的化学物质。如植物激素。 , 重力作用于根冠细胞造粉质体，根的伸长区产生反应，由IAA传递信息 导致根生长的向重性。 , 玉米，向日葵等根尖遭受干旱后迅速合成ABA，然后通过导管向地上部 运输(此时木质部伤流液中ABA可增加25-30倍)，到达叶细胞的质外体， 气孔保卫细胞Ca2+浓度增加，质膜去极化产生K+外流和苹果酸下降， 保卫细胞失水，引起气孔关闭。 , 局部伤害引起周身性反应的信息传导途经，当番茄植株一个叶片遭虫咬 伤后，会诱导叶片ABA增加，pin基因活化，产生蛋白酶抑制物(proteinase inhibitor，PI)以阻碍病原或害虫进一步侵害。 , 将受害叶的细胞壁水解产物(主要是寡聚糖)加到叶片中，又可模拟伤害反 应诱导PIs的产生，认为寡聚糖是由受伤叶片释放并经维管束转移，继而 诱导能使PIs基因活化的化学信号物质。 2. 物理信号(physical signal): 指细胞感受到刺激后产生的能够起传递信息作用的物理因子。如电、光、磁场、水力学等可以在生物细胞间或其内起信号分子的作用。 当用一个微电极插入一个未受刺激的细胞内时，可以记录到细胞内有电位差，称为静息电位。一般细胞中这种电位差为内负外正，在-100mv—50mV之间，即正常细胞一般都处于极化状态。当给予一个刺激时，会引起静息电位发生一定程度的去极化(内正外负)的电位变化，称为动作电位(指细胞和组织中发生的相对于空间和时间的快速变化的一类生物电位,离子流膜两边分布导致电位变化)。 电信号是生物体内最重要的物理信号。它主要指细胞膜静息电位改变时所引起动作电位的定向传播，它在植物环境刺激—细胞反应系统中起重要作用，是植物体长距离传递信息的一种重要方式。 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 电信号传递特点:对高敏感的植物，外界刺激无需达到伤害程度即可产生动作电波(action potential AP);中等敏感植物在伤害刺激条件下产生变异电波(variation potential VP);不敏感的植物只引起不可传递的局部电位变化，表明植物都有经逆境或剧烈刺激激活的潜在兴奋性。 物理信号传递特点: , 导致化学信号产生:与动物相似，植物的电波也是质膜极化及透性变化 的结果，而且拌有化学信号的产生(如乙酰胆碱生成)。 , 电信号传递途径:植物电波长途传递途径是微管束，短距离传递则通过 共质体和质外体。 , 电信号作用:各种电波传递都可以产生生理效应。 例举: Wildon等(1992)用番茄实验发现，当番茄幼苗的子叶受到机械伤害时，导致动作电位从子叶中缓慢发出并且进入第一片真叶，在此与诱导PI(proteinase inhibitor)的所有反应相联系。所以伤害刺激产生的电信号，在番茄幼苗中传递，可诱导pin基因活化产生蛋白酶抑制物。证明了电信号可引起包括基因转录在内深刻生理生化变化。 植物对水力学信号(hydraulic signal)即水压的变化很敏感。玉米叶片木质部压力的微小变化就能迅速影响叶片气孔开度，即压力降低时气孔开放，压力提高时气孔关闭。 (二 )跨膜信号转换机制 胞外信号少部分可以直接跨过细胞膜系统引起生理反应;多数则与膜系统上的受体(receptor)识别后，通过膜上信号转换系统，转变为胞内信号，才能调节细胞代谢及生理功能。能与受体结合的特殊信号物质称配体(Ligang)。受体可以是蛋白质，也可以是一个酶系。 1.受体与信号的感受: , 物理信号以辐射、电磁场、力等方式使细胞内外受体直接感受刺激，引obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 起受体结构变化而活化。如光受体(色素蛋白)通过多肽链上结合生色 团(色素)来接受光信号，生色团接受光信号后发生构象变化，进而带 动多肽链变构使光受体活化。 , 化学信号可通过配基(或激子)与受体在空间结构上的互补性、化学键 (氢键、离子键)、作用力(范德华力)、生物分子(酶蛋白与底物或抑 制剂、抗原和抗体)间的亲和力等方式与受体发生特异性结合或相互作 用，使受体感知信号并引发受体分子构象变化而将其激活。 受体存在部位 存在于细胞表面的细胞质膜受体，大多信号分子不能通过膜，信号分子通过与细胞表面受体结合，经过跨膜信号转换，将胞外信号转至胞内。 细胞内受体是存在于细胞内即细胞质或亚细胞组分(细胞核)上受体。 目前研究较多的如下三类受体: 植物激素受体(hormone receptor)能与激素特异结合并引起特殊生理效应的物质，一般是属于蛋白质。 光信号受体: 对红光和远红光敏感的光敏色素(Pr、Pfr),对蓝光敏感的蓝光受体(隐花色素)、对紫外光敏感的紫外光受体(UV-B)。 感病诱导因子受体。 (分属化学、物理、生物受体) 激素受体，生长素受体在细胞中的存在位置有多种说法，但主要有两种: 一种存在于质膜上 它能促进细胞壁松驰，是酸生长理论。 另一种存在于细胞质(或细胞核)中 它能促进核酸和蛋白质的合成，是基因活化学说。 光受体:光敏色素。迄今，光敏色素已被纯化，基因克隆。其中从藓类(Certodon Rurpureus)中克隆了一个新的光敏色素基因，并证实该光敏色素N端(生色团所在)作为光受体分子，C端则在光刺激后产生蛋白激酶活性。 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 2.G蛋白(G protein)转换: 又称信号转换蛋白或偶联蛋白。全称为GTP结合调节蛋白(GTP binding regulatory protein),此类蛋白由于其生理活性有赖于三磷酸鸟苷(GTP)的结合以及具有GTP水解的活性而得名. 二十世纪70年代初在动物细胞中发现了G蛋白的存在，进而证明了G蛋白是细胞膜受体与其所调节的相应生理过程之间的主要信号转导者。 G蛋白的发现是生物学一大成就。吉尔曼(Gilman)与罗德贝尔(Rodbell)因此获得1994年诺贝尔医学生理奖。 G蛋白一般分为两大类: 异源三体G蛋白:三种亚基(,、,、,)构成 小 G 蛋 白:一个亚基的单体 G蛋白功能体现，是靠与GTP的结合或水解产生的变构作用完成。当G蛋白与受体结合而激活时，它就同时结合上GTP，继而触发效应器，把外界信号转换为胞内信号;而当GTP水解为GDP后，G蛋白就回到原初构象，失去转换的功能。 植物G蛋白的研究始于上世纪80年代，现已证明，G蛋白在高等植物中普遍存在，并初步证明它通过光、植物激素对植物产生生理效应，如跨膜离子运输、气孔运动、植物形态建成等生理活动的细胞信号转导过程中有重要调节作用。为信号转导主要方式。 由于细胞表面受体对信号的感知和转换过程不同，细胞中质膜上的三种类型的受体 (A)G蛋白偶连受体 活化时G蛋白连接受体传递信息到G蛋白质，其上有GTP。 GTP复合体中的α-亚基能与β、γ亚基分开，进入细胞质激活其他酶。调节生理过程:种子萌发、细胞分裂、气孔开闭、光形态建成、对逆境反应等。 (B)酶偶连受体 受体通常是蛋白激酶，与信号结合后，随受体活化，内部分子磷酸化，传递信息，将胞外信号传至胞内。 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. (C) 离子通道偶连受体 受体可能本身是细胞表面重要的通道。接受信号时，通道开放。使细胞内离子浓度发生变化，这一离子浓度变化便是第二信使(Ca2+信号)。 二、胞内信号传导 来自胞外的各种物理或化学刺激信号作为细胞信号转导过程中的初级信号或第一信使，把由胞外刺激信号激活或抑制的(或引起改变的)，具有生理调节活性的细胞内因子，称为细胞信号转导过程中的次级信号或第二信使(second messenger)。 目前研究较多的胞内信号有三类: 钙信号系统: Ca++ CaM 肌醇磷脂信号系统: 以肌醇磷脂代谢为基础，IP3(三磷酸肌醇)、DAG(二酰甘油) 环核苷酸信号系统: cAMP 1. 钙信号系统 (1)衡量钙信使的 标准 excel标准偏差excel标准偏差函数exl标准差函数国标检验抽样标准表免费下载红头文件格式标准下载 , 细胞质的Ca2+水平必须能对来自环境与临近细胞的刺激有所反应;而且 Ca2+水平的变化要早于该生理反应; , 在缺乏正常的外界刺激时，通过诱导细胞质Ca2+浓度的变化，能引起该 生理反应; , 细胞必须具有感受Ca2+浓度变化，并将这些变化转变为某种生理反应的 机制; , 阻拦第二信使感知系统的运行，必定会阻拦对外界刺激的生理反应。 (2)细胞游离Ca2+分布与转移 通常细胞中钙(总钙)以结合态和自由离子态(Ca2+)两种形式存在。一般植物细胞在未受到刺激时(静息态)其细胞质Ca2+水平都相当低，约为10-6,10-7mol/L,而细胞外(细胞壁及间隙)可达10-3mol/L。 细胞壁是细胞最大的钙库，细胞器如线粒体，叶绿体，微体，液胞，内质网obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 等的Ca2+浓度也是细胞质的几百倍到几千倍。因次，在细胞质中与胞外或胞内钙库(某些细胞器)之间存在一个Ca2+浓度梯度。 若细胞质内Ca2+浓度过高会产生毒害，将会同磷酸反应形成沉淀而扰乱以磷酸为基础的能量代谢。因此，植物细胞必须具有从胞质中主动派出过量Ca2+的机制，而使细胞内Ca2+浓度维持在低水平。 一般来说，细胞内Ca2+浓度十分稳定，即使有微小拨动，也是短暂的，故称为钙稳态(calcium homeostasis)。液泡大量积累Ca2+并形成不溶性钙盐(草酸钙，苹果酸钙等)可看作是从细胞溶质中清除过量Ca2+的手段之一。 (3)钙信使作用的分子基础 Ca2+受体及钙调蛋白 Ca2+作为第二信使，是如何起到调节酶和细胞功能作用的呢,植物细胞的钙信号受体蛋白之一是钙结合蛋白(CaBP)，它与Ca2+有很高的亲和力和专一性，即使当细胞液中Mg+浓度大于Ca2+浓度1000倍时，仍优先与Ca2+结合。钙结合蛋白中分布最广，目前最普遍的了解最多的是钙调素(Calmodulin，CaM)。 钙调素生物化学 CaM是由19种氨基酸组成的一种耐热，酸性(等电点3.9-4.3)小分子可溶性球蛋白，95?加热3min不变性，十分稳定。148个氨基酸组成的单链小分子(17KD,19KD)。植物CaM的分子量，氨基酸组成和基本理化性质都与动物CaM相似，不同的是植物CaM多一个半胱氨酸残基。另外，动物CaM含两个酪氨酸，而植物只有一个。 Ca与钙调素的作用机理 在一个体系内，Ca2+作为信号是基于其浓度的变化。通常细胞溶质Ca2+水平通常低于10-6mol.L-1以下，而胞外则可达10-3mol.L-1。胞内Ca2+浓度变化可通过引起膜透性变化，具体通过开启/关闭膜上的Ca2+通道而引起。 , 当某刺激(信号分子)到达细胞时，质膜上Ca2+通道打开，对Ca2+通透 性瞬间增加， Ca2+流入细胞质，使细胞质Ca2+浓度增加，到达一定阈obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 值(高于10-6mol.L-1)。 , 细胞质中，Ca与CaM结合，形成复合体，这种激活状Ca*CaM即可进 而与它所调节的靶酶结合而激活靶酶(Ca-CaM.E) ，完成特定的生理生化 过程。 , 之后Ca2+与CaM分离，CaM与靶酶复合体解离，CaM与靶酶均回到 非活性状态;这时细胞质中的Ca2+通过膜上Ca2+-ATP酶，将Ca2+从 细胞质泵入钙库。细胞质Ca2+降低到与CaM结合阈值以下，细胞质中 Ca2+又回到静息态水平。 目前有十多种酶受Ca2+-CaM调节。如蛋白激酶，NAD激酶，H+-ATPase等和光敏素为受体的光信号转换中起重要的调节作用。 通过细胞质内Ca2+浓度变化而把细胞外的信息传递给细胞内各相关过程的功能，即为Ca2+的信号功能。 实验证明，几乎所有的胞外刺激信号(光照，温度，重力，触摸等物理刺激和各种植物激素，病原菌诱导因子等化学物质)都可引起胞内游离Ca2+浓度的变化，而这种Ca2+浓度变化的时间，幅度，频率，区域化分布等却不尽相同，而表现生理效应也不同。 如GA的作用表现一种适度而持久(min到h量级)的Ca2+增加，缓慢回落;IAA的作用常表现为Ca2+振荡(即出现重复的Ca2+峰)。不同刺激信号的特异性可能就是靠Ca2+浓度变化的不同形式而体现的。 Ca的作用方式: (1)直接与靶酶结合，诱导靶酶的活性构象而调节它们的活性，如NAD 激酶、 Ca2+ -ATP激酶等; (2)通过活化依赖 Ca2+ ?CaM的蛋白激酶，将靶酶磷酸化，影响其活性，如磷酸化酶、H+-ATP酶等。 2. 肌醇磷脂信号系统 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 肌醇磷脂是一类由磷脂酸与肌醇结合的脂质化合物。肌醇分子六碳环上的羟基被不同数目的磷酸酯化主要以三种形式存在于植物质膜上: 磷脂酰肌醇(PI) 磷脂酰肌醇-4-磷酸(PIP) 磷脂酰肌醇-4，5-二磷酸(PIP2)。 以肌醇磷脂代谢为基础的细胞信号系统，是外界信号被膜受体接受后，以G蛋白为中介，由质膜中的磷脂酶 C(PLC，是一类磷脂酰键的水解酶,在生物生命活动中起着第二信使的重要作用 )水解PIP2，生成肌醇-1，4，5-三磷酸(IP3)和二酰甘油(DAG)两种信号分子。 PLC作用产生的信号物质IP3和DAG作用。 IP3通过调节Ca2+浓度调节酶活性; DAG则通过激活蛋白激酶C(PKC，依赖Ca2+/钙调蛋白的蛋白激酶 )传递信息。由膜质成份肌醇磷脂产生的IP3和DAG称为双信号。 IP3作用: 具有使贮钙体液泡释放Ca2+的功能。一方面， Ca2+控制PLC的底物专一性，决定IP3和DAG信号分子的生成;另一方面，IP3信使的量反过来又影响Ca2+的产生，表明植物细胞中钙信使与肌醇磷脂信号系统的共辖性质。已有证据表明，IP3/ Ca2+信号系统在干旱和ABA引起的蚕豆，豌豆气孔开闭，雨树小叶膨压运动，植物对病原微生物侵染等环境刺激-快速反应中起信号转导作用。 DAG作用: 作为信号分子蛋白激酶C(PKC)的受体。PKC的活性依赖于Ca2+和磷脂，它催化蛋白质的磷酸化。在正常情况下，质膜上不存在自由的DAG，它只是在细胞受到外界刺激时肌醇磷脂水解才产生。在有DAG和Ca2+时，磷脂与PKC分子相结合，使PKC激活，从而对某些底物蛋白活酶类进行磷酸化，导致细胞反应。 3. 环核苷酸信号系统 在动物细胞中早已证明了cAMP是重要的第二信使物质。一般认为，当某些胞外刺激信号作用于动物细胞时，质膜上的受体通过G蛋白介导，促进或抑制膜内侧的腺苷酸环化酶，从而调节胞质内cAMP水平，cAMP作为第二信使进obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 而调节细胞的生理生化活动。 自70年代人们就开始研究植物细胞内的cAMP，但至今进展缓慢，仍未证明它在植物细胞中的普遍存在以及是否也具有作为胞内第二信使的作用。在叶绿体光诱导花色素苷合成过程中，cAMP参与受体G蛋白之后的下游信号转导过程. 1994年Bowler等报道了利用分子生物学和细胞显微注射方法，以光敏色素A缺失的番茄突变系为材料，研究了完整叶绿体中多种光系统色素蛋白质及酶基因表达所必须的光信号传递过程。 三、蛋白质的磷酸化和去磷酸化 植物体内许多功能蛋白转录后需要共价修饰才能发挥其生理功能，由蛋白激酶(protein kinase)/蛋白磷酸脂酶(protein phosphatases)所催化的磷酸化/去磷酸化就是进行共价修饰的一个重要过程。 这些酶可以作为胞内第二信使作用的靶酶，即胞内信号通过调节磷酸化或去磷酸化而进一步进行转导信号。如载体磷酸化和脱磷酸化与功能的改变。蛋白激酶是一类催化蛋白质产生磷酸化的酶，可对其底物蛋白质特定的氨基酸残基进行磷酸化修饰，引起相应的生理反应。从最初信号刺激、G蛋白转化、第二信使、激活相应蛋白到促使蛋白磷酸化等一系列反应的逐级放大。完成信号转导过程。第二信使调节激酶: , 蛋白激酶的底物可以是酶，因此它们既可以直接通过对酶的磷酸化修饰 来改变酶的活性,完成特定的生理过程。 , 蛋白激酶也可以是转录因子，可通过修饰转录因子(transcription factors) 而激活或抑制基因的表达，从而使细胞对外来信号作出相应的反应。 1(蛋白激酶 在植物中，目前已知的蛋白激酶至少有30多种，其中最重要的蛋白激酶有两类: obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 一类钙依赖型蛋白激酶( Ca2+ -dependent protein kinase，CDPK)或称钙调素样蛋白激酶(Calmodulin-like domain protein kinase，CDPK)。它最初从大豆培养细胞中分离鉴定出，现已证明是植物细胞中一个蛋白激酶家族，仅从拟南芥中克隆的CDPK基因就有10种左右。 此酶由508个氨基酸组成，在其氨基端有一激酶活性区域，而在其羧基端有一类似于CaM的调节结构区域(其中有4个Ca2+结合位点)，在两者之间存在由31个氨基酸组成的自身抑制区域(autoinhibitory domain)。当Ca2+与CDPK结合时，其自身抑制区域对激酶活性的抑制被解除，CDPK被活化。这类激酶活性只依赖于Ca2+而不依赖于CaM。 在植物中，目前已知的蛋白激酶至少有30多种，它们的作用也表现在多个方面，包括向光性、抗寒、抗病、根部的向地性、光合作用、自交不亲和性以及细胞分裂等。 除CDPK类蛋白激酶外，还有与植物光敏色素密切相关的Ca2+依赖性蛋白激酶、同时受Ca2+和CaM调节的蛋白激酶。 研究得较多的是最初从大豆中得到的依赖于钙离子的蛋白激酶，CDPK有一与钙结合位点。现已知的可被CDPK磷酸化的作用靶物质(或底物分子)有: 细胞骨架成分、膜运输成分、质膜上的质子ATP酶等。如从燕麦中分离出与质膜成分相结合的CDPK成分可将质膜上的质子ATP酶磷酸化，从而调节跨膜离子的运输。 另一是类受体蛋白激酶， 在动物细胞表面有一类受体称为受体蛋白激酶，后来在植物中发现与之同源的基因，由于基因产物的受体功能未能得到证实，故将它们称为类受体蛋白激酶(receptor-like protein kinase, RLK)。这些蛋白激酶可能与调节花粉自交不亲和、病源信号转导以及生长发育有关过程(受体蛋白激酶具有特异性)。 2(蛋白磷酸酶 蛋白磷酸脂酶逆转磷酸化作用，是终止信号或一种逆向调节，与蛋白激酶理论上有同等重要意义。现在植物中鉴定出几种不同的蛋白质磷酸酯酶。如obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 豌豆、胡萝卜中的Type1蛋白磷酸酯酶、豌豆、胡萝卜、小麦中的Type2A和Type2C蛋白磷酸酯酶等。 有研究表明，Type1蛋白磷酸酯酶可能与植物细胞的有丝分裂过程的调控有关。在豌豆保卫细胞中存在一种依赖钙离子的蛋白磷酸酯酶，它与K+的转移和气孔的开闭有关。 虽然磷酸化和去磷酸化的过程本身是单一的反应，但不同蛋白质磷酸化和去磷酸化的结果却是不同的，很可能与实现细胞中各种不同刺激信号的转导过程有关。实际上，正是蛋白质磷酸化的可逆性为细胞的信息传递提供了一种开关作用;在有外来信号刺激的情况下，通过脱磷酸化而失活能够再将之关闭，这就使得细胞能够有效而经济的调控对内外信息的反应。 四、细胞的生理反应 细胞反应是细胞信号转导的最后一步，外界信号刺激一般都能引起相应的细胞反应。 所有外界刺激都能引起相应的细胞反应，细胞对不同外界信号的生理反应也不同，如外界刺激可引起多种生理变化: 1.跨膜离子流动，离子运输、气孔调节等; 2.细胞骨架的变化，质体运动、有机物及信息在胞内运输和传递; 3.细胞内代谢途径的调控，有机物合成分解及转化等; 4.细胞内相应基因的表达。 信号转导中，根据植物从感受刺激到表现出相应生理反应的时间，植物的生理反应可分为长期效益和短期效益。 , 短期生理效应:如气孔反应、含羞草感振反应、转板藻的叶绿体运动、 棚田效应等。 , 长期生理效应:二对生长发育的调控的信号转导来说，大部分为长期生 理效应，如光对种子萌发、春化作用、光周期效应对开花调控等。持续 的信号作用代谢调节，对基因转录的调节，使植物内部产生深刻变化。 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 细胞生理反应的行为变化类型包括: 细胞代谢:使细胞摄入并代谢营养物质，提供细胞生命活动所需要能量。 细胞分裂:使与DNA复制相关基因表达，调节细胞周期，使细胞进入分裂和增殖阶段。 细胞分化:使细胞内遗传程序有选择表达，细胞最终不可逆分化成特定功能成熟细胞。 细胞功能:如激素、蛋白、核苷酸等，使细胞能够进行正常的代谢活动。 细胞死亡:在局部范围内和一定数量上发生细胞程序性死亡，以维护多细胞生物的整体利益或减轻不良环境的危害。 一些常见的植物信号转导事例 生理现象 信号 受体或感受部位 相应生理生化反应 植物向光性反蓝光 向光素 茎受光侧IAA高于背光应 侧而生长速率低 光诱导种子萌红光远红光 光敏色素 红光促进种子萌发远红发 光抑制 光诱导气孔运蓝光，绿光 蓝光受体，玉米蓝光促进气孔开发绿光动 黄素 抑制 干旱气孔运动 干旱 细胞壁或膜 ABA合成与气孔关闭 根向地性生长 重力 根冠筑细胞淀粉IAA向地与背地分布不 体 同，生长速率不同 含羞草震感运机械刺激，电感受细胞的膜 离子跨膜运输，叶枕膨动 波 压变化，小叶运动 光周期与开花 光周期 光敏色素和隐花相关开花基因表达，花 色素 芽分化 低温诱导开花 低温 茎尖分生组织 相关开花基因表达，花 芽分化 果实成熟 乙烯 乙烯受体 纤维素及果胶酶基因表 达，物质转化，膜透性obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No. 增加 植物抗病反应 病原体的激发激发子受体 植保素，病原相关蛋白 子 豆科植物根瘤 根瘤的结瘤因凝集素 促进根皮层根瘤形成 子 obstacles, correcting misunderstandings, advocate good cadre style. 2, adhere to the scientific decision-making and democratic decision-making decision-making law. Correctly handle to ensure that government decrees and based on the actual creative work, giving full play to subjective initiative, put an end to implement the conference meeting, to document the implementation of documents etc. Improve the scientific and democratic decision-making mechanism, improve and implement the decision to solicit the views of experts, the public hearing system. Improve the important decision of risk assessment mechanism and legal review mechanism. 3, to promote team unity. Uphold and improve the system of annual party book talk heart to heart, mind and Party members, Party members to exchange each other not less than 2 times. The implementation of sound team democratic life system, consolidate and enhance the party's mass line educational practice will promote the achievement of the topic of democratic life, criticism and self-criticism normalization, a long-term. Adhere to the party on the basis of the principle of unity, enhance internal communication, strengthen internal supervision, self Sleep maintenance team collective image and authority. (three) to ensure the execution is not strong or disguised to resist government decrees issues. Individual Bureau leadership team members lack of learning political theory system, not from the objective reality, not with the party's interests above all the principles to deal with personal problems. In the specific work safety supervision in the execution is not strong, in the implementation of the decision to deploy the discount, make choices, engage in work, the implementation of the agreement, uncomfortable is not performed, open or disguised to resist, the lack of subject consciousness of the party organization, the lack of resist in society all kinds of bad habits and unwholesome tendencies of courage Gas and courage. Such as the liability insurance of safe production work, individual leaders that have someone arrested, and no funding, not in the global chess perspective to the countryside to carry out daily inspection, found none of my business acceptance of the resumption of production in an armchair, with a pen, indifferent. Corrective measures: (LED Leadership: Luo Mingjun, rectification time: before September 25th, insist for a long time) 1, to strengthen the sense of responsibility, work to solve the lack of decent occupation explain away, conduct problems. To establish the overall concept, eliminate departmentalism. Strict assessment and accountability, to solve the spiritual slack, nianqingpazhong, status quo, To establish and perfect the muddle along and other issues. Bureau staff conduct work regulations, standardize the behavior of personnel. 2, strengthen the responsibility system. One is the in-depth study and implement the "Hunan provincial Party and government leading cadres Interim Provisions on the work safety of a pair of > (Hunan Office issued 2013 No.