水杨酸对大豆耐铝性的调控机制
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摘要
铝(Aluminum,Al)作为酸性土壤上最主要的非生物胁迫因子严重影响了植物生长和作物产量。对大多数植物而言,土壤中微摩尔浓度的Al即能明显地抑制根伸长。根尖是Al毒害最敏感的部位,Al离子与根尖相互作用,在几分钟之内就能导致植物根伸长受到显著抑制,根尖膨大,严重阻碍植物对养分和水分的吸收。Al诱导大豆根系分泌柠檬酸被认为是其最重要的耐Al机制之一。然而,调控柠檬酸分泌的过程和Al抑制根伸长的机理目前尚不十分清楚。
水杨酸(Salicylic acid,SA)作为关键的信号分子参与调控植物体内的多种生理功能,如种子萌发、幼苗生长和各种生物和非生物胁迫响应。在过去几年的研究中,SA在植物抗病方面起着重要的信号调节作用,如诱导植物抗病基因(PR)的表达、激活过敏反应(HR)和系统获得性抗性(SAR)。而最近的研究表明,SA在植物抗非生物胁迫(紫外线辐射、低温、热激、干旱、盐胁迫和重金属毒害等逆境)方面也发挥着至关重要的作用。有证据显示,SA不仅与柠檬酸代谢关键酶-顺乌头酸酶(ACO)和柠檬酸转运体MATE家族密切相关,而且SA介导ROS的产生以及诱导相关基因的表达很可能与调控植物的防御反应有关。因此,本文以大豆耐Al品种吉育70(Glycine max L. cv Jiy u 70)为试验材料,采用液体培养方法,分别从柠檬酸分泌、抗氧化系统、转录因子(Gm-WRKY58和Gm-NPR1)和ABC转运体(Gm-STAR1和Gm-PDR12)基因表达的角度深入探讨外源SA对大豆耐铝性的调控机制,得到如下主要研究结果。
1.水杨酸对大豆耐铝性的调控作用
外源SA缓解Al对根伸长的抑制作用,且存在浓度依赖及“低促高抑”效应,低浓度(10μmmol/L)SA处理能显著缓解Al对根伸长的抑制,降低Al在根尖中的积累。外源供给SA能快速被根尖吸收,同时大豆内源SA能响应外源Al胁迫与SA处理,内源自由态SA含量在Al胁迫处理4 h达到最高,结合态SA(SAG)含量随Al处理时间的延长而显著增加。SA生物合成抑制剂多效挫(PAC)能抑制Al诱导自由态SA的产生,但对SAG没有影响。这说明外源SA在缓解大豆Al毒害过程中起作用,SA很可能作为Al胁迫的早期信号分子调控大豆耐Al性。
2.水杨酸对铝胁迫诱导大豆根系柠檬酸分泌的调控作用
Al胁迫下植物根系分泌有机酸是植物重要的抗Al机制之一。目前关于SA是如何促进Al胁迫下柠檬酸分泌的调控机制尚不清楚。本文试图从顺乌头酸酶(ACO)和柠檬酸转运体(MATE)的角度来阐明SA与Al胁迫诱导大豆根系柠檬酸分泌的关系。Al胁迫条件下,外源SA显著提高了Al诱导大豆根系柠檬酸的分泌,PAC不但降低了Al诱导柠檬酸的分泌,且能够抑制自由态SA的产生,表明内源SA对柠檬酸分泌起调控作用。Al和SA共处理与单独Al处理相比进一步降低线粒体ACO的活性,但各处理在不同时间点(2、4、9、12h)对ACO基因的表达差异并不显著,说明ACO很可能在蛋白水平上受到调控;Al和SA共处理与单独Al处理相比提高了大豆根尖柠檬酸转运体Gm-MATE1基因的转录表达,然而Al胁迫条件下外源SA对大豆根尖Gm-MATE1基因的表达和Al诱导柠檬酸分泌之间并不呈线性关系,说明Gm-MATE1基因的表达很可能是柠檬酸分泌的一个前提。以上试验结果表明,Al胁迫条件下SA在柠檬酸的代谢和运输过程中起着至关重要的作用。
3.水杨酸对铝诱导大豆根尖氧化胁迫的调控作用
Al能诱导活性氧(ROS)的产生,而ROS的产生和清除可以通过抗氧化系统相关酶来控制。近年来的诸多研究结果表明,SA在调控受胁迫的植物体内氧化还原内稳态上起了十分重要的作用。为了进一步探讨SA的效果,本文对抗氧化系统的相关生理指标进行了测定。结果表明,Al胁迫条件下,通过内源SA的快速应答,可引起大豆根尖H_2O_2水平发生显著变化。外源SA能迅速激活H_2O_2且在6h达到峰值,随着处理时间的延长(9、12h),SA反而降低了Al诱导H_2O_2的积累。说明外源SA增强大豆抗Al胁迫能力可能与过氧化氢(H_2O_2)的变化存在一定的联系,且Al胁迫条件下SA很可能激活了H_2O_2的信号转导途径。SA通过调控H_2O_2代谢相关酶如SOD、CAT、APX的活性,使大豆根尖H_2O_2含量维持在一定水平,进而缓解Al诱导的氧化胁迫伤害。外源SA降低了Al诱导大豆根尖质膜过氧化产物丙二醛(MDA)和超氧阴离子(O_2~(·–))的累积。上述试验结果表明,SA也可以通过对抗氧化系统的调控进一步缓解Al毒害。
4.水杨酸对铝诱导转录因子Gm-WRKY58、Gm-NPR1及ABC转运体Gm-STAR1、Gm-PDR12基因表达的影响
已有证据表明,在植物响应胁迫的过程中SA能有效地调控转录因子WRKY和NPR1基因的表达。ABC转运体基因编码膜结合蛋白,是目前已知功能最广泛的超蛋白家族之一。ABC转运体不仅涉及激素、脂类、金属、次生代谢物以及异生物质的运输,而且对离子通道的调控也起着十分重要的作用。本试验结合前期大豆在铝胁迫下基因芯片的分析结果,利用实时荧光定量PCR技术进行分析,结果表明,单独Al处理均显著增加了大豆根尖Gm-WRKY58、Gm-NPR1、Gm-STAR1、Gm-PDR12基因的相对表达水平;而与单独Al处理相比,外源SA分别不同程度地增加了Al诱导的这些基因在根尖的相对表达。表明,转录因子Gm-WRKY58、Gm-NPR1以及ABC转运体Gm-STAR1、Gm-PDR12可能在SA调控的大豆根尖Al胁迫响应过程中起到一定的作用。
Approximately 40% of the word’s potentially arable soils are acidic and negatively impacted by the Aluminum (Al). It is a universal problem that Al toxicity has therefore been represented one of the major constraint on plant growth and crop production in acid soil worldwide. Al-induced citrate exudation has been identified as an important Al-resistance mechanism in plants. Salicylic acid (SA), considered as a crucial signal molecule, plays an important role in regulating the plant disease resistance. It has been proposed that SA can involve in modulation of acclimation responses to abiotic stresses such as low temperature, heat shock, drought, salinity, heavy metal toxicity. Thus, this study was to research the changes of endogenous SA under Al stress; clarify SA involved in the process of Al induced-citrate exudation from soybean roots; explore the effect of exogenous SA on the antioxidant system under Al stress.
Root elongation is a rapid and efficient parameter to assessment of Al resistance. The obtained results indicated that Al-inhibited root elongation response to different concentrations of applied SA differs and the obvious relief in Al toxicity caused by exogenous 10μM SA. The proper concentration of SA is an important factor in determining plant response to Al stress. 10μM SA decreased the Al accumulation in soybean root tips. Endogenous SA content of soybean root tips have a rapidly response to Al stress and exogenous SA treatment. The data was shown that external supply of SA to roots is quickly taken up by the root. the SA content of the root tips increased up 4 h, and following decrease with the elongation of Al treatment duration. SAG accumulated gradually throughout the treatment period. In order to further elucidate the role of SA under Al stress, an inhibitor of SA biosynthesis (PAC) was used in this study. PAC application slightly but significantly reduced endogenous free SA content in root tips of Al and SA-treated plants. However the effect on SAG was less clear. Al and SA together treatment increased the amount of Al-induced citrate secretion from soybean roots, further decreased aconitase (ACO) activity and enhanced the transcriptional expression of Gm-MATE1 compared to the sole Al treatment. From the above results, it is suggested that SA production might act as an early signal to regulate the metabolism and transportation of citrate in soybean roots.
SA modulated the H_2O_2 level in root tips of soybean by regulating the activity of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) which could take response to Al stress. Exogenous SA decreased the Al-induced enhancement of MDA in soybean root tips. Taken together, the possible of mechanisms of SA under Al stress are reducing the ACO activity and increasing Gm-MATE1 gene expression, leading to enhance citrate efflux, and then, alleviated the Al-induced inhibition of root elongation and decreased the Al accumulation in soybean root tips. SA may also regulate the antioxidant system to alleviate Al toxicity.
There is evidence that SA can be effective in regulating the transcription factor WRKY and NPR1 in the process of plants respond to stress.ABC transporter genes encoding membrane-bound protein, is the most widely known function of the ultra-protein family.ABC transporters are not only involved in hormones, lipids, metals, secondary metabolites and xenobiotic transport, but also play an important role in regulation of ion channels. In this study, according to microarray results, using real-time quantitative PCR technology, analysis the genes expression of transcription factors Gm-WRKY58 and Gm- NPR1 as well as ABC transporters Gm-STAR1 and Gm- PDR12 under Al and/or SA treatments. The results show that, Al induced an increase genes relative expression of Gm-WRKY58、Gm-NPR1、Gm-SATR1 and Gm-PDR12 rapidly;adding exogenous SA enhanced the Al induced these genes expression. The obtained results indicated that the genes expression of Gm-WRKY58 and Gm-NPR1 as well as Gm-SATR1 and Gm-PDR12 response to Al stress, may play a crucial role in SA regulate Al resistance in soybean roots.
水杨酸(Salicylic acid,SA)作为关键的信号分子参与调控植物体内的多种生理功能,如种子萌发、幼苗生长和各种生物和非生物胁迫响应。在过去几年的研究中,SA在植物抗病方面起着重要的信号调节作用,如诱导植物抗病基因(PR)的表达、激活过敏反应(HR)和系统获得性抗性(SAR)。而最近的研究表明,SA在植物抗非生物胁迫(紫外线辐射、低温、热激、干旱、盐胁迫和重金属毒害等逆境)方面也发挥着至关重要的作用。有证据显示,SA不仅与柠檬酸代谢关键酶-顺乌头酸酶(ACO)和柠檬酸转运体MATE家族密切相关,而且SA介导ROS的产生以及诱导相关基因的表达很可能与调控植物的防御反应有关。因此,本文以大豆耐Al品种吉育70(Glycine max L. cv Jiy u 70)为试验材料,采用液体培养方法,分别从柠檬酸分泌、抗氧化系统、转录因子(Gm-WRKY58和Gm-NPR1)和ABC转运体(Gm-STAR1和Gm-PDR12)基因表达的角度深入探讨外源SA对大豆耐铝性的调控机制,得到如下主要研究结果。
1.水杨酸对大豆耐铝性的调控作用
外源SA缓解Al对根伸长的抑制作用,且存在浓度依赖及“低促高抑”效应,低浓度(10μmmol/L)SA处理能显著缓解Al对根伸长的抑制,降低Al在根尖中的积累。外源供给SA能快速被根尖吸收,同时大豆内源SA能响应外源Al胁迫与SA处理,内源自由态SA含量在Al胁迫处理4 h达到最高,结合态SA(SAG)含量随Al处理时间的延长而显著增加。SA生物合成抑制剂多效挫(PAC)能抑制Al诱导自由态SA的产生,但对SAG没有影响。这说明外源SA在缓解大豆Al毒害过程中起作用,SA很可能作为Al胁迫的早期信号分子调控大豆耐Al性。
2.水杨酸对铝胁迫诱导大豆根系柠檬酸分泌的调控作用
Al胁迫下植物根系分泌有机酸是植物重要的抗Al机制之一。目前关于SA是如何促进Al胁迫下柠檬酸分泌的调控机制尚不清楚。本文试图从顺乌头酸酶(ACO)和柠檬酸转运体(MATE)的角度来阐明SA与Al胁迫诱导大豆根系柠檬酸分泌的关系。Al胁迫条件下,外源SA显著提高了Al诱导大豆根系柠檬酸的分泌,PAC不但降低了Al诱导柠檬酸的分泌,且能够抑制自由态SA的产生,表明内源SA对柠檬酸分泌起调控作用。Al和SA共处理与单独Al处理相比进一步降低线粒体ACO的活性,但各处理在不同时间点(2、4、9、12h)对ACO基因的表达差异并不显著,说明ACO很可能在蛋白水平上受到调控;Al和SA共处理与单独Al处理相比提高了大豆根尖柠檬酸转运体Gm-MATE1基因的转录表达,然而Al胁迫条件下外源SA对大豆根尖Gm-MATE1基因的表达和Al诱导柠檬酸分泌之间并不呈线性关系,说明Gm-MATE1基因的表达很可能是柠檬酸分泌的一个前提。以上试验结果表明,Al胁迫条件下SA在柠檬酸的代谢和运输过程中起着至关重要的作用。
3.水杨酸对铝诱导大豆根尖氧化胁迫的调控作用
Al能诱导活性氧(ROS)的产生,而ROS的产生和清除可以通过抗氧化系统相关酶来控制。近年来的诸多研究结果表明,SA在调控受胁迫的植物体内氧化还原内稳态上起了十分重要的作用。为了进一步探讨SA的效果,本文对抗氧化系统的相关生理指标进行了测定。结果表明,Al胁迫条件下,通过内源SA的快速应答,可引起大豆根尖H_2O_2水平发生显著变化。外源SA能迅速激活H_2O_2且在6h达到峰值,随着处理时间的延长(9、12h),SA反而降低了Al诱导H_2O_2的积累。说明外源SA增强大豆抗Al胁迫能力可能与过氧化氢(H_2O_2)的变化存在一定的联系,且Al胁迫条件下SA很可能激活了H_2O_2的信号转导途径。SA通过调控H_2O_2代谢相关酶如SOD、CAT、APX的活性,使大豆根尖H_2O_2含量维持在一定水平,进而缓解Al诱导的氧化胁迫伤害。外源SA降低了Al诱导大豆根尖质膜过氧化产物丙二醛(MDA)和超氧阴离子(O_2~(·–))的累积。上述试验结果表明,SA也可以通过对抗氧化系统的调控进一步缓解Al毒害。
4.水杨酸对铝诱导转录因子Gm-WRKY58、Gm-NPR1及ABC转运体Gm-STAR1、Gm-PDR12基因表达的影响
已有证据表明,在植物响应胁迫的过程中SA能有效地调控转录因子WRKY和NPR1基因的表达。ABC转运体基因编码膜结合蛋白,是目前已知功能最广泛的超蛋白家族之一。ABC转运体不仅涉及激素、脂类、金属、次生代谢物以及异生物质的运输,而且对离子通道的调控也起着十分重要的作用。本试验结合前期大豆在铝胁迫下基因芯片的分析结果,利用实时荧光定量PCR技术进行分析,结果表明,单独Al处理均显著增加了大豆根尖Gm-WRKY58、Gm-NPR1、Gm-STAR1、Gm-PDR12基因的相对表达水平;而与单独Al处理相比,外源SA分别不同程度地增加了Al诱导的这些基因在根尖的相对表达。表明,转录因子Gm-WRKY58、Gm-NPR1以及ABC转运体Gm-STAR1、Gm-PDR12可能在SA调控的大豆根尖Al胁迫响应过程中起到一定的作用。
Approximately 40% of the word’s potentially arable soils are acidic and negatively impacted by the Aluminum (Al). It is a universal problem that Al toxicity has therefore been represented one of the major constraint on plant growth and crop production in acid soil worldwide. Al-induced citrate exudation has been identified as an important Al-resistance mechanism in plants. Salicylic acid (SA), considered as a crucial signal molecule, plays an important role in regulating the plant disease resistance. It has been proposed that SA can involve in modulation of acclimation responses to abiotic stresses such as low temperature, heat shock, drought, salinity, heavy metal toxicity. Thus, this study was to research the changes of endogenous SA under Al stress; clarify SA involved in the process of Al induced-citrate exudation from soybean roots; explore the effect of exogenous SA on the antioxidant system under Al stress.
Root elongation is a rapid and efficient parameter to assessment of Al resistance. The obtained results indicated that Al-inhibited root elongation response to different concentrations of applied SA differs and the obvious relief in Al toxicity caused by exogenous 10μM SA. The proper concentration of SA is an important factor in determining plant response to Al stress. 10μM SA decreased the Al accumulation in soybean root tips. Endogenous SA content of soybean root tips have a rapidly response to Al stress and exogenous SA treatment. The data was shown that external supply of SA to roots is quickly taken up by the root. the SA content of the root tips increased up 4 h, and following decrease with the elongation of Al treatment duration. SAG accumulated gradually throughout the treatment period. In order to further elucidate the role of SA under Al stress, an inhibitor of SA biosynthesis (PAC) was used in this study. PAC application slightly but significantly reduced endogenous free SA content in root tips of Al and SA-treated plants. However the effect on SAG was less clear. Al and SA together treatment increased the amount of Al-induced citrate secretion from soybean roots, further decreased aconitase (ACO) activity and enhanced the transcriptional expression of Gm-MATE1 compared to the sole Al treatment. From the above results, it is suggested that SA production might act as an early signal to regulate the metabolism and transportation of citrate in soybean roots.
SA modulated the H_2O_2 level in root tips of soybean by regulating the activity of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT) and ascorbate peroxidase (APX) which could take response to Al stress. Exogenous SA decreased the Al-induced enhancement of MDA in soybean root tips. Taken together, the possible of mechanisms of SA under Al stress are reducing the ACO activity and increasing Gm-MATE1 gene expression, leading to enhance citrate efflux, and then, alleviated the Al-induced inhibition of root elongation and decreased the Al accumulation in soybean root tips. SA may also regulate the antioxidant system to alleviate Al toxicity.
There is evidence that SA can be effective in regulating the transcription factor WRKY and NPR1 in the process of plants respond to stress.ABC transporter genes encoding membrane-bound protein, is the most widely known function of the ultra-protein family.ABC transporters are not only involved in hormones, lipids, metals, secondary metabolites and xenobiotic transport, but also play an important role in regulation of ion channels. In this study, according to microarray results, using real-time quantitative PCR technology, analysis the genes expression of transcription factors Gm-WRKY58 and Gm- NPR1 as well as ABC transporters Gm-STAR1 and Gm- PDR12 under Al and/or SA treatments. The results show that, Al induced an increase genes relative expression of Gm-WRKY58、Gm-NPR1、Gm-SATR1 and Gm-PDR12 rapidly;adding exogenous SA enhanced the Al induced these genes expression. The obtained results indicated that the genes expression of Gm-WRKY58 and Gm-NPR1 as well as Gm-SATR1 and Gm-PDR12 response to Al stress, may play a crucial role in SA regulate Al resistance in soybean roots.
引文
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