生育后期养分胁迫对水稻衰老进程影响的蛋白质组学研究
摘要
水稻既是世界重要的粮食作物之一,又是基因组学研究的模式材料,在生产实践和科学研究中都占有极其重要的地位。杂交水稻具有适应性广、多抗、高产、优质等特点,但部分杂交稻组合存在早衰现象,水稻早衰不仅影响后期干物质的生产和积累,也会影响前期积累的干物质转化为产量的实现,影响籽粒灌浆和干物质的运输与分配,最终阻碍产量潜力的发挥。杂交水稻早衰是一个相对复杂的生理生化过程,除受遗传因素影响外,环境条件也是水稻发生早衰的重要诱导因子。
本研究采用双向电泳分离技术、蛋白质质谱分析技术以及生物信息学等手段,对养分胁迫下易早衰杂交早稻威优916灌浆期根、叶、叶鞘及籽粒蛋白质组的差异表达进行分析,探讨杂交水稻早衰机理及在不同组织器官中的代谢特征,是学科前沿与实际应用的有机结合,在科研和生产实践中都具有重要的意义。
首先,不同处理下易早衰杂交早稻威优916根、叶、叶鞘及籽粒的四个时期可溶性蛋白经双向电泳分离,构建了相应器官的蛋白质组表达图谱,并对发生了差异表达的121个目的蛋白质点进行了MALDI-TOF-MS/MS分析和数据库检索鉴定,共有89个蛋白质功能得到鉴定,成功率为73.55%。这些被鉴定的蛋白质根据其功能可以分为6个不同类别,其中绝大数为光合作用、抗性及逆境信号传递、特定器官生长发育相关蛋白质。
生育后期持续的养分胁迫,诱导了威优916剑叶中的部分蛋白质发生了差异表达,其中绝大部分为光合作用、信号传导及胁迫抗性相关蛋白质。与光合作用相关蛋白质如转酮醇酶、核酮糖二磷酸羧化酶/加氧酶大亚基、核酮糖磷酸盐羧化酶主链蛋白前体等在整个养分胁迫期间表达量均是大幅下降,表明叶片的光合能力显著下降,光合产物减少。信号传导及胁迫抗性相关蛋白质包括丝氨酸蛋白激酶、半胱氨酸合成酶及S-腺苷甲硫氨酸合成酶等,养分胁迫诱导了叶片中大量该类蛋白质的积累,表明叶片能够感知并传递外界胁迫信号,使其能够迅速启动逆境防御系统,最大限度的减少逆境伤害,而类胚素蛋白的高表达和过氧化氢酶类的低表达,会导致叶片中大量H202积累,从而破坏叶片的正常生理活动。
叶鞘中受养分胁迫影响的蛋白质大部分为信号传导及胁迫抗性相关蛋白质,其次为光合作用相关蛋白质。信号传导及胁迫抗性相关蛋白质包括铁蛋白、分泌型过氧化物酶及谷胱甘肽硫转移酶等,养分胁迫导致叶鞘中这些保护酶类在处理两周后的表达量逐渐下调,而充分的养分供应则使叶鞘中保护酶表达量持续上调,意味着养分胁迫导致叶鞘的抗逆性减弱,而逆境响应蛋白质V型ATP酶和乙醇脱氢酶表达量上调表达,叶鞘感知胁迫加剧并传递信号。养分胁迫导致叶鞘中叶绿素合成受阻,直接参与光合作用蛋白质表达量下调,叶鞘光合能力下降。
因养分胁迫而发生差异表达的根系蛋白质中,绝大部分为根系生长发育和逆境胁迫响应相关蛋白质。养分胁迫前中期诱导了威优916根系中包括阿拉伯呋喃糖苷酶、RGP蛋白、几丁质酶等根系生长类蛋白质的上调表达,从而促进了根系的快速分化和生长,是植物根系养分缺乏环境下的一种自我调节机制,通过增大根系吸收面积来提高吸收效率。逆境胁迫响应相关蛋白质包括甘油-3-磷酸脱氢酶、过氧化物酶植物凝集素等,这些蛋白质均为胁迫应激反应类蛋白质,这些蛋白质在根系中的上调表达,表明根系感知胁迫反应剧烈。
养分胁迫对水稻的影响最终都会通过籽粒这个特定的器官表现出来,养分胁迫导致籽粒的结实率和千粒重显著下降。通过对籽粒的蛋白质组表达变化分析发现,养分胁迫诱导了威优916籽粒中的部分蛋白质发生了差异表达,其中绝大多数为籽粒充实发育相关蛋白质,如谷蛋白、WD40结构域蛋白、胚蛋白等,这类蛋白质在快速灌浆的前中期表达量下调,导致籽粒灌浆和胚发育不良。而籽粒中热激蛋白质Hsp70、半胱氨酸合成酶、分支酸变位酶等逆境抗性及信号传递蛋白质的差异表达,表明籽粒作为最终的“库”器官同样能够感知胁迫信号,但保护类蛋白质的下调表达也意味着植株的抗性减弱。
本研究利用蛋白质组学方法来揭示养分胁迫对易早衰水稻不同器官组织的衰老进程影响,鉴定了一些发生了差异表达的蛋白质,通过对其功能及表达量变化的分析,为更好的理解水稻早衰生理代谢机理提供依据。
Rice is one of the most important staple food crops for human in the world, which plays important role in the agricultural production and scientific research. Besides, it is a model monocot for genome study. Hybrid rice has some merits such as broad adaptability, high resistance, high yield and quality. However, the phenomenon of premature senescence widely appears in the combination of inter-subspecific hybridization, which not only impacts the assimilation and accumulation of dry matter during late growth stage, but also affects the grains filling, dry matter transportation and distribution, hence decreases the potential of high yield. As a very complicated genetic physiological process, premature senescence can be induced by genetic and environmental factors.
Hybrid rice Weiyou916was found to be easily induced for premature senescence under nutrient stress. Therefore, classical proteomic approaches including two-dimensional electrophoresis (2-DE), matrix assisted laser desorption/ionization time of flight mass spectrometry (MALDI-TOF/MS) and bioinformatics technique were used to analyze the differential expression proteomics of roots, leaves, leaf sheathes and grains of Weiyou916. in order to find out the premature senescence mechanism of hybrid rice and metabolism character in different tissues under nutrient stress. It would be the bridge between research prospect and practical application, and it would provide important value in the field of scientific research and agricultural production.
The2-DE technology was used to determine the tissue proteome maps of roots, leaves, leaf sheathes and grains of Weiyou916at four different stages, respectively. Totally,121proteins were analyzed by MALDI-TOF/MS and database searching, among which89proteins were identified. The success ratio of proteins identification was73.55%. The identified proteins could be classified into6functional categories. Most of these proteins involved in photosynthesis, resistance, stress signal transportation, and tissue development.
The proteins in flag leaves of Weiyou916performed differential expression under nutrient stress at late development stage. Most of these proteins involved in photosynthesis, signal transduction and resistance. The expression amount of proteins associated with photosynthesis such as transketolase, ribulose bisphosphate carboxylase and precursor of ribulose bisphosphate carboxylase main chain were greatly decreased under nutrient stress, which resulted in low photosynthetic rate and low accumulation of dry matter in flag leaves. Lots of proteins involved in signal transduction and resistance such as serine/threonine protein kinases, cysteine synthase and S-adenosylmethionine synthetase in Weiyou916flag leaves were induced and accumulated under nutrient stress, indicating that rice plant could immediately trigger off the resistance system by receptors and signal transduction, and that the great amount of expression on these proteins reduced the damage of stress. Nevertheless, the accumulation of H2O2in flag leaves were induced by high amount of germin-like protein and low amount of catalase, which disturbed the normal physiological activities of leaves.
Most of proteins induced by nutrient stress in leaf sheathes were involved in signal transduction and stress resistance, and minority of proteins were related to the photosynthesis. The expression amount of proteins related to signal transduction and stress resistance including ferritin, secretory peroxidases and glutathione S-transferase were decreased under nutrient stress after two weeks, which indicated low resistance, but those enzymes in sheathes under full nutrient supply increased in expression. Vacuolar ATPase and alcohol dehydrogenase involved in stress response increased in protein expression, which showed that leaf sheathes could sense and transfer the environment stress information. The biosynthesis of chlorophyll and photosynthesis proteins in leaf sheathes were inhibited by nutrient stress, hence decreasing photosynthesis of leaf sheathes.
Most of protiens with differential expression due to nutrient stress in roots were related to roots growth, development and stress resistance. Proteins involved in growth such as arabinofuranosidase. reversibly glycosylated polypeptide and chitinase have shown drastically increase in response to nutrient stress, which promoted root growth. It could attribute to the fact that roots had a self-regulation mechanism and enhanced the nutrient uptake by increasing the radicular absorption area under insufficient nutrient condition. GPDH. peroxidase and jasmonate-induced lectin were the proteins which associated stress resistance and stress response, and the increase of these proteins indicated that roots had a sensitive-receptor.
Finally, nutrient stress resulted in low seed setting rate and1000-grain weight. The analysis of grain proteome by proteomics technology showed that nutrient stress resulted in differential expression in some proteins in grains. Among those proteins, some are associated with grain filling and development, such as glutelin, WD40domain and germin-like protein presented low expression, which resulted in low grain filling rate and germ dysplasia. However, the Hsp70protein, cysteine synthases and chorismate mutase involved in stress resistance and signal transduction showed low expression, which indicated that grains could receive the stress signal and attenuation resistance in plant.
In this study, proteome was used to determine the effect of nutrient stress on rice tissues, valuable functional and abundant information of some differential expression proteins was analyzed and acquired. The results would contribute to a deep understanding on the mechanism of early-ageing hybrid rice.
本研究采用双向电泳分离技术、蛋白质质谱分析技术以及生物信息学等手段,对养分胁迫下易早衰杂交早稻威优916灌浆期根、叶、叶鞘及籽粒蛋白质组的差异表达进行分析,探讨杂交水稻早衰机理及在不同组织器官中的代谢特征,是学科前沿与实际应用的有机结合,在科研和生产实践中都具有重要的意义。
首先,不同处理下易早衰杂交早稻威优916根、叶、叶鞘及籽粒的四个时期可溶性蛋白经双向电泳分离,构建了相应器官的蛋白质组表达图谱,并对发生了差异表达的121个目的蛋白质点进行了MALDI-TOF-MS/MS分析和数据库检索鉴定,共有89个蛋白质功能得到鉴定,成功率为73.55%。这些被鉴定的蛋白质根据其功能可以分为6个不同类别,其中绝大数为光合作用、抗性及逆境信号传递、特定器官生长发育相关蛋白质。
生育后期持续的养分胁迫,诱导了威优916剑叶中的部分蛋白质发生了差异表达,其中绝大部分为光合作用、信号传导及胁迫抗性相关蛋白质。与光合作用相关蛋白质如转酮醇酶、核酮糖二磷酸羧化酶/加氧酶大亚基、核酮糖磷酸盐羧化酶主链蛋白前体等在整个养分胁迫期间表达量均是大幅下降,表明叶片的光合能力显著下降,光合产物减少。信号传导及胁迫抗性相关蛋白质包括丝氨酸蛋白激酶、半胱氨酸合成酶及S-腺苷甲硫氨酸合成酶等,养分胁迫诱导了叶片中大量该类蛋白质的积累,表明叶片能够感知并传递外界胁迫信号,使其能够迅速启动逆境防御系统,最大限度的减少逆境伤害,而类胚素蛋白的高表达和过氧化氢酶类的低表达,会导致叶片中大量H202积累,从而破坏叶片的正常生理活动。
叶鞘中受养分胁迫影响的蛋白质大部分为信号传导及胁迫抗性相关蛋白质,其次为光合作用相关蛋白质。信号传导及胁迫抗性相关蛋白质包括铁蛋白、分泌型过氧化物酶及谷胱甘肽硫转移酶等,养分胁迫导致叶鞘中这些保护酶类在处理两周后的表达量逐渐下调,而充分的养分供应则使叶鞘中保护酶表达量持续上调,意味着养分胁迫导致叶鞘的抗逆性减弱,而逆境响应蛋白质V型ATP酶和乙醇脱氢酶表达量上调表达,叶鞘感知胁迫加剧并传递信号。养分胁迫导致叶鞘中叶绿素合成受阻,直接参与光合作用蛋白质表达量下调,叶鞘光合能力下降。
因养分胁迫而发生差异表达的根系蛋白质中,绝大部分为根系生长发育和逆境胁迫响应相关蛋白质。养分胁迫前中期诱导了威优916根系中包括阿拉伯呋喃糖苷酶、RGP蛋白、几丁质酶等根系生长类蛋白质的上调表达,从而促进了根系的快速分化和生长,是植物根系养分缺乏环境下的一种自我调节机制,通过增大根系吸收面积来提高吸收效率。逆境胁迫响应相关蛋白质包括甘油-3-磷酸脱氢酶、过氧化物酶植物凝集素等,这些蛋白质均为胁迫应激反应类蛋白质,这些蛋白质在根系中的上调表达,表明根系感知胁迫反应剧烈。
养分胁迫对水稻的影响最终都会通过籽粒这个特定的器官表现出来,养分胁迫导致籽粒的结实率和千粒重显著下降。通过对籽粒的蛋白质组表达变化分析发现,养分胁迫诱导了威优916籽粒中的部分蛋白质发生了差异表达,其中绝大多数为籽粒充实发育相关蛋白质,如谷蛋白、WD40结构域蛋白、胚蛋白等,这类蛋白质在快速灌浆的前中期表达量下调,导致籽粒灌浆和胚发育不良。而籽粒中热激蛋白质Hsp70、半胱氨酸合成酶、分支酸变位酶等逆境抗性及信号传递蛋白质的差异表达,表明籽粒作为最终的“库”器官同样能够感知胁迫信号,但保护类蛋白质的下调表达也意味着植株的抗性减弱。
本研究利用蛋白质组学方法来揭示养分胁迫对易早衰水稻不同器官组织的衰老进程影响,鉴定了一些发生了差异表达的蛋白质,通过对其功能及表达量变化的分析,为更好的理解水稻早衰生理代谢机理提供依据。
Rice is one of the most important staple food crops for human in the world, which plays important role in the agricultural production and scientific research. Besides, it is a model monocot for genome study. Hybrid rice has some merits such as broad adaptability, high resistance, high yield and quality. However, the phenomenon of premature senescence widely appears in the combination of inter-subspecific hybridization, which not only impacts the assimilation and accumulation of dry matter during late growth stage, but also affects the grains filling, dry matter transportation and distribution, hence decreases the potential of high yield. As a very complicated genetic physiological process, premature senescence can be induced by genetic and environmental factors.
Hybrid rice Weiyou916was found to be easily induced for premature senescence under nutrient stress. Therefore, classical proteomic approaches including two-dimensional electrophoresis (2-DE), matrix assisted laser desorption/ionization time of flight mass spectrometry (MALDI-TOF/MS) and bioinformatics technique were used to analyze the differential expression proteomics of roots, leaves, leaf sheathes and grains of Weiyou916. in order to find out the premature senescence mechanism of hybrid rice and metabolism character in different tissues under nutrient stress. It would be the bridge between research prospect and practical application, and it would provide important value in the field of scientific research and agricultural production.
The2-DE technology was used to determine the tissue proteome maps of roots, leaves, leaf sheathes and grains of Weiyou916at four different stages, respectively. Totally,121proteins were analyzed by MALDI-TOF/MS and database searching, among which89proteins were identified. The success ratio of proteins identification was73.55%. The identified proteins could be classified into6functional categories. Most of these proteins involved in photosynthesis, resistance, stress signal transportation, and tissue development.
The proteins in flag leaves of Weiyou916performed differential expression under nutrient stress at late development stage. Most of these proteins involved in photosynthesis, signal transduction and resistance. The expression amount of proteins associated with photosynthesis such as transketolase, ribulose bisphosphate carboxylase and precursor of ribulose bisphosphate carboxylase main chain were greatly decreased under nutrient stress, which resulted in low photosynthetic rate and low accumulation of dry matter in flag leaves. Lots of proteins involved in signal transduction and resistance such as serine/threonine protein kinases, cysteine synthase and S-adenosylmethionine synthetase in Weiyou916flag leaves were induced and accumulated under nutrient stress, indicating that rice plant could immediately trigger off the resistance system by receptors and signal transduction, and that the great amount of expression on these proteins reduced the damage of stress. Nevertheless, the accumulation of H2O2in flag leaves were induced by high amount of germin-like protein and low amount of catalase, which disturbed the normal physiological activities of leaves.
Most of proteins induced by nutrient stress in leaf sheathes were involved in signal transduction and stress resistance, and minority of proteins were related to the photosynthesis. The expression amount of proteins related to signal transduction and stress resistance including ferritin, secretory peroxidases and glutathione S-transferase were decreased under nutrient stress after two weeks, which indicated low resistance, but those enzymes in sheathes under full nutrient supply increased in expression. Vacuolar ATPase and alcohol dehydrogenase involved in stress response increased in protein expression, which showed that leaf sheathes could sense and transfer the environment stress information. The biosynthesis of chlorophyll and photosynthesis proteins in leaf sheathes were inhibited by nutrient stress, hence decreasing photosynthesis of leaf sheathes.
Most of protiens with differential expression due to nutrient stress in roots were related to roots growth, development and stress resistance. Proteins involved in growth such as arabinofuranosidase. reversibly glycosylated polypeptide and chitinase have shown drastically increase in response to nutrient stress, which promoted root growth. It could attribute to the fact that roots had a self-regulation mechanism and enhanced the nutrient uptake by increasing the radicular absorption area under insufficient nutrient condition. GPDH. peroxidase and jasmonate-induced lectin were the proteins which associated stress resistance and stress response, and the increase of these proteins indicated that roots had a sensitive-receptor.
Finally, nutrient stress resulted in low seed setting rate and1000-grain weight. The analysis of grain proteome by proteomics technology showed that nutrient stress resulted in differential expression in some proteins in grains. Among those proteins, some are associated with grain filling and development, such as glutelin, WD40domain and germin-like protein presented low expression, which resulted in low grain filling rate and germ dysplasia. However, the Hsp70protein, cysteine synthases and chorismate mutase involved in stress resistance and signal transduction showed low expression, which indicated that grains could receive the stress signal and attenuation resistance in plant.
In this study, proteome was used to determine the effect of nutrient stress on rice tissues, valuable functional and abundant information of some differential expression proteins was analyzed and acquired. The results would contribute to a deep understanding on the mechanism of early-ageing hybrid rice.
引文
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