结缕草温度胁迫的生理响应及其分子机制研究
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摘要
温度作为生物机能的一种动力，影响植物的蒸腾、水势、吸收和新陈代谢，以及几乎所有的酶促反应、休眠和生长发育。草坪草的生长温度域很窄，冷季型草坪草的适宜温度为15～24℃，暖季型草坪草的适宜温度为26～32℃，超过适宜温度，草坪草的正常生长发育和健康就会受到影响。因此，草坪草抗寒耐热性研究一直是热点问题。国内对草坪草抗性的研究多局限于生理生化方面的变化机制，对其分子机理研究甚少。本试验以暖季型草坪草为材料，系统研究草坪草对温度胁迫的生理响应及其分子适应机制。
以南方最广泛种植的两种暖季型草坪草——沟叶结缕草(Zoysiagrass matrella(L．)merr．)和日本结缕草(Zoysiagrass japonica Steud)为材料，测定其高低温胁迫下主要抗性生理指标的变化。结果表明：两种结缕草在高温和低温胁迫下，叶片可溶性糖、可溶性蛋白质和脯氨酸等渗透调节物质的含量都有不同程度增加，其中沟叶结缕草不管是高温下还是低温下其叶片可溶性糖和脯氨酸含量的增幅明显大于日本结缕草。超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)是细胞膜的保护酶系统，具有清除活性氧物质的功能。试验结果表明，低温和高温下，两种结缕草叶片SOD、POD和CAT三种酶活性都有不同程度增强，其中沟叶结缕草在逆境温度下其SOD与CAT活性增强均大于日本结缕草，日本结缕草则表现为逆境下POD活性增强大于沟叶结缕草。MDA是膜脂过氧化的主要产物之一，它的积累是活性氧毒害的表现，本试验结果表明，高温下两种结缕草MDA含量增加不显著，但在低温下，其含量都极显著增加，其中日本结缕草MDA含量的增加幅度大于沟叶结缕草。光合色素的含量不仅影响草坪草的色泽，而且影响其光合作用。在本试验中，高低温胁迫下两种结缕草叶片的叶绿素a(Chla)、叶绿素b(Chlb)、类胡萝卜素(Car)含量以及Chla／Chlb的比值均不同程度下降，尤其是日本结缕草。植物激素在植物生长过程中起重要作用。本试验结果表明，两种结缕草在高低温胁迫下叶片抑制类激素ABA含量都增加，40℃高温胁迫下两种草坪草ABA含量增幅均大于0℃或10℃低温胁迫，同样高温或低温胁迫下沟叶结缕草的ABA含量增幅又大于日本结缕草；高温胁迫下，两种结缕草叶片促进类激素都相对保持较高含量，在低温胁迫时促进类激素含量则有较大幅度降低。从各内源激素间的平衡来看，在高低温胁迫下，两种结缕草叶片内抑制类激素与促进类激素总量的比值升高。
综合各抗性生理指标值的变化，我们认为两种结缕草抗高温性能优于抗低温；同样的温度胁迫下，沟叶结缕草的抗逆性又强于日本结缕草。
在前期温度胁迫的抗性生理指标测定与分析的基础上，选择抗性表现更佳的沟叶结缕草为材料，利用双向电泳技术(two-dimensional electrophoresis，2-DE)、质谱技术(mass spectrometry，MS)并结合生物信息学的方法，分别对其在低温胁迫和高温胁迫条件下的差异蛋白质组学进行了研究。结果发现高低温下共有35个蛋白质点在不同处理间存在显著的差异表达模式。35个蛋白质点通过Q-TOF MS分析后有26个蛋白获得了有效的肽指纹图谱和功能预测。根据其功能，将它们归纳为五个类群，即：信号转导相关蛋白、代谢相关蛋白、防卫相关蛋白、蛋白质合成与降解相关蛋白、细胞转运能力和胚胎发育相关蛋白。其中，在低温胁迫下，质谱分析共鉴定出20个蛋白点，有4个点是同一个蛋白(Rubisco大亚基)，这些差异表达的蛋白质一方面加强了胞内信号转导、活性氧的清除、蛋白质防御以提高沟叶结缕草的抗逆性，另一方面由于一些蛋白质丰度的下调表达使得低温下植物体呼吸作用表现为先增强后减弱、光合作用下降、蛋白质降解、细胞的转运能力减弱，不利植物的生长发育；在高温胁迫下，质谱分析共鉴定出19个蛋白点，3个点是同一个蛋白(Rubisco大亚基)，这些蛋白质主要加强了信号识别、跨膜信号转导、光合作用、呼吸作用、次生代谢作用、活性氧的清除、蛋白质防御及蛋白质的合成等方面，极大地提高了沟叶结缕草的高温抗逆性。此外，在本试验高低温处理中，均鉴定出ZLL／PNH同源蛋白相对丰度的下调表达，且均随着时间的延长，下调幅度增大，由此说明，不管是在高温胁迫下还是在低温胁迫下均不利沟叶结缕草主茎分生组织的发育，不利于维持主茎的正常生理机能。
进一步运用抑制消减杂交(SSH)方法构建沟叶结缕草对高温(40℃)处理响应的正向差减文库，通过reverse northern杂交方法筛选文库，挑选出高温胁迫处理下35个差异表达基因，对这些基因进行测序，将测序获得的35条原始序列去除由SSH引入的接头引物序列后与基因组数据库(NCBI)进行比对，获得26个基因功能已知的差异表达的基因。这些已知基因分别参与植物代谢、基因表达调控和信号转导、植物抗逆性反应以及蛋白质的合成等过程。试验中鉴定出与代谢有关的NADP-苹果酸酶、与蛋白质合成有关的核糖体蛋白以及具有清除活性氧功能的过氧化氢酶等基因的加强表达，这一结果与高温胁迫下沟叶结缕草的差异蛋白质组学研究的结果相一致，进一步证明了高温胁迫下沟叶结缕草通过加强代谢作用，增强蛋白质的合成作用以及提高活性氧的清除能力来提高其高温抗逆性。文中还讨论了SSH分析结果与蛋白质组学分析结果不一致的可能原因。
Temperature as a drive force of biological functions affects plant's transpiration,absorption,water potential,nearly all enzymatic reations,dormancy and growth.Thetemperature range of growth in turfgrass is narrow.The suitable temperature range of growthfor cold-season turfgrass is 15～24℃,while it is 26～32℃for warm-season turfgrass.Whenit exceeds the suitable temperature range,the natual growth and health of turfgrass will beinfluenced.Studying on the cold resistance and heat tolerance of turfgrass is a hot issue.Somestudies on the physiobiochemical mechanism of the resistance in turfgrass have been carriedout,however few information is available in the molecular mechanism of its resistance inChina.The study is to detect the physiological responses and molecular adaptationmechanism of warm-season turfgrass to temperature stress.
Two warm-type turfgrass species,the most widely planted,such as Zoysia matrella (L.)merr.and Zoysia japonica steud.,were used for the experiment to detect the physiologicalresponses to low and high temperature stress.The results showed that the contents of solublesugar,soluble protein,proline,were all increased,but the increasing changes in the contentsof soluble sugar and proline were higher in Z.matrella than in Z.japonica under the sametemperature stress.Superoxide dismutase (SOD),peroxidase (POD)and catalase (CAT),which function is clearing active oxygen,are the system of protective enzymes in cellmembrane.The study showed that the activities of SOD,POD and CAT were all builded up inthe two turfgrass species under the low and high temperature stresses.But the increased extent ofSOD and CAT in Z.matrella was much higher than that in Z.japonica,however Z.japonica showedhigher activity of POD than Z.matrella under the temperature stress.Malondialdehyde (MDA)isone major production of membrane lipid peroxidation.Its accumulation is physiologicalindicator of active oxygen toxicity.It was found that the content of MDA was insignificantlyincreased under the high temperature stress,but it was significantly enhanced under the lowtemperature stress in the two turfgrass species.The increasing extent of MDA content wasmuch higher in Z.matrella than that in Z.japonica under the low temperature stress.Photosynthetic pigments not only affect the colour and luster of turfgrass but also affect itsphotosynthesis.It was also found that low and high temperature treatment caused decreases in carotenoid (Car)、chlorophyll a (Chla)and chlorophyll b (Chlb)contents and changes inChla/Chlb.However,the two turfgrass species responded to the temperature stressdifferently.The decreasing extent of the pigments was higher in Z.japonica under thetemperature treatment.Endogenous hormones play an important role on the plant growth.Thestudy showed that the content of ABA was increased when exposed to temperature treatment,and the increased extent was much higher under 40℃treatment than under 0℃or10℃treatment in the two turfgrass species.The increased degree of ABA in Z.matrella wasmuch higher than that in Z.japonica under the same temperature treatment.It maitainedrelatively high contents of promoting hormones in the two turf grass species under the hightemperature treatment,but reverse was true in the low temperature treatment.As far as thebalance of endogenous hormones was concerned,the ratio of inhibiting hormones topromoting hormones was increased in the two turfgrass species under the temperaturetreatment.
According to the results obtained from analyzing the resistance physiological indexes,itindicated that the two turfgrass species performed stronger resistibility under the hightemperature stress than under the low temperatue stress and Z.matrella showed the strongerresistance than Z.japonica under the same temperature stress.
In this article,the warm season turfgrass,Z.matrella,which showed stronger resistancein the experiment of resistance physiology,was studied using two-dimensional gelelectrophoresis(2-DE),mass spectrometry(MS),and bioinformatics analyses to explore thedifferentially expressed proteins of the turfgrass under low temperature and high temperaturetreatments.The result indicated that there were 35 potein spots which showed differentexpression partterns after the the different treatments.Of these,26 proteins were successfullyidentified with the aid of Q-TOF MS and bioinformatics.These proteins were involved insignal transduction,metabolism,stress resistance,protein synthesis and degradation,celltransportation capacity and embryo development.There were 20 proteins identified by Q-TOFMS,of which 4 proteins were ribulose bisphosphate carboxylase large subunit,under the lowtemperature treatment.On the one hand,some of these proteins strengthened signaltransduction,active oxygen scavenging and protein defencing,which were attributed to havethe strong resistant ability in Z.matrella,on the other hand,some other proteins,which was down-regulated,the respiration was strengthen firstly and then weaken,photosythesis wasdecreased,protein degraded,and transportatio ability of cell organ reduced in Z.matrella,inturn caused the plant growth trouble.There were 19 proteins successfully identified byQ-TOF MS,of which 3 proteins were ribulose bisphosphate carboxylase large subunit,underthe high temperature treatment.These proteins strengthened signal identification,transmembrane signal transduction,photosythesis,respiration,secondary metabolism,activeoxygen scavenging,protein defencing and protein synthesis,and led to the strong resistibilityof Z.matrella under the high temperature treatment.Besides that,the relative abundance ofZLL/PNH homologous protein decreased and the decreasing extent was increased with thetime prolonged under the low temperature treatment.The similar tendency was also foundunder the high temperature stress.These suggested that it was unfavorable for the main stemmeristematic tissue development and growth either under the high temperatue stress or underthe low temperatue stress.
Further,an investigation was conducted to detect the related gene expression profiles in Z.matrella exposed to high temperature stress using suppression subtractive hybridization(SSH)method.A total of 35 ESTs from SSH-cDNA library were sequenced,and all of thesequences of inserted fragments were analyzed with the aid of bioinformatics.It obtained 26different functionally known genes.These genes were sorted into four groups such asmetabolism,gene expression regulation and signal transduction,stress resistance and proteinsynthesis.The expressions in genes of NADP-dependent malic protein,ribosomal protein S 1and catalase,which respectively involved in metabolism,protein synthesis and active oxygenscavenging,were enhanced under the high temperature stress.This result is consistent to thoseof differential proteomics research,and in turn further proved that the resistance wasstrengthened by improving metabolism,protein synthesis and the ability to scavenge activeoxygen in Z.matrella under high temperature stress.Finally,the author also discussed why alot of strengthen expression genes,which were identified in the experiment,were differentfrom outcomes of differential proteomics research in this paper.
以南方最广泛种植的两种暖季型草坪草——沟叶结缕草(Zoysiagrass matrella(L．)merr．)和日本结缕草(Zoysiagrass japonica Steud)为材料，测定其高低温胁迫下主要抗性生理指标的变化。结果表明：两种结缕草在高温和低温胁迫下，叶片可溶性糖、可溶性蛋白质和脯氨酸等渗透调节物质的含量都有不同程度增加，其中沟叶结缕草不管是高温下还是低温下其叶片可溶性糖和脯氨酸含量的增幅明显大于日本结缕草。超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)是细胞膜的保护酶系统，具有清除活性氧物质的功能。试验结果表明，低温和高温下，两种结缕草叶片SOD、POD和CAT三种酶活性都有不同程度增强，其中沟叶结缕草在逆境温度下其SOD与CAT活性增强均大于日本结缕草，日本结缕草则表现为逆境下POD活性增强大于沟叶结缕草。MDA是膜脂过氧化的主要产物之一，它的积累是活性氧毒害的表现，本试验结果表明，高温下两种结缕草MDA含量增加不显著，但在低温下，其含量都极显著增加，其中日本结缕草MDA含量的增加幅度大于沟叶结缕草。光合色素的含量不仅影响草坪草的色泽，而且影响其光合作用。在本试验中，高低温胁迫下两种结缕草叶片的叶绿素a(Chla)、叶绿素b(Chlb)、类胡萝卜素(Car)含量以及Chla／Chlb的比值均不同程度下降，尤其是日本结缕草。植物激素在植物生长过程中起重要作用。本试验结果表明，两种结缕草在高低温胁迫下叶片抑制类激素ABA含量都增加，40℃高温胁迫下两种草坪草ABA含量增幅均大于0℃或10℃低温胁迫，同样高温或低温胁迫下沟叶结缕草的ABA含量增幅又大于日本结缕草；高温胁迫下，两种结缕草叶片促进类激素都相对保持较高含量，在低温胁迫时促进类激素含量则有较大幅度降低。从各内源激素间的平衡来看，在高低温胁迫下，两种结缕草叶片内抑制类激素与促进类激素总量的比值升高。
综合各抗性生理指标值的变化，我们认为两种结缕草抗高温性能优于抗低温；同样的温度胁迫下，沟叶结缕草的抗逆性又强于日本结缕草。
在前期温度胁迫的抗性生理指标测定与分析的基础上，选择抗性表现更佳的沟叶结缕草为材料，利用双向电泳技术(two-dimensional electrophoresis，2-DE)、质谱技术(mass spectrometry，MS)并结合生物信息学的方法，分别对其在低温胁迫和高温胁迫条件下的差异蛋白质组学进行了研究。结果发现高低温下共有35个蛋白质点在不同处理间存在显著的差异表达模式。35个蛋白质点通过Q-TOF MS分析后有26个蛋白获得了有效的肽指纹图谱和功能预测。根据其功能，将它们归纳为五个类群，即：信号转导相关蛋白、代谢相关蛋白、防卫相关蛋白、蛋白质合成与降解相关蛋白、细胞转运能力和胚胎发育相关蛋白。其中，在低温胁迫下，质谱分析共鉴定出20个蛋白点，有4个点是同一个蛋白(Rubisco大亚基)，这些差异表达的蛋白质一方面加强了胞内信号转导、活性氧的清除、蛋白质防御以提高沟叶结缕草的抗逆性，另一方面由于一些蛋白质丰度的下调表达使得低温下植物体呼吸作用表现为先增强后减弱、光合作用下降、蛋白质降解、细胞的转运能力减弱，不利植物的生长发育；在高温胁迫下，质谱分析共鉴定出19个蛋白点，3个点是同一个蛋白(Rubisco大亚基)，这些蛋白质主要加强了信号识别、跨膜信号转导、光合作用、呼吸作用、次生代谢作用、活性氧的清除、蛋白质防御及蛋白质的合成等方面，极大地提高了沟叶结缕草的高温抗逆性。此外，在本试验高低温处理中，均鉴定出ZLL／PNH同源蛋白相对丰度的下调表达，且均随着时间的延长，下调幅度增大，由此说明，不管是在高温胁迫下还是在低温胁迫下均不利沟叶结缕草主茎分生组织的发育，不利于维持主茎的正常生理机能。
进一步运用抑制消减杂交(SSH)方法构建沟叶结缕草对高温(40℃)处理响应的正向差减文库，通过reverse northern杂交方法筛选文库，挑选出高温胁迫处理下35个差异表达基因，对这些基因进行测序，将测序获得的35条原始序列去除由SSH引入的接头引物序列后与基因组数据库(NCBI)进行比对，获得26个基因功能已知的差异表达的基因。这些已知基因分别参与植物代谢、基因表达调控和信号转导、植物抗逆性反应以及蛋白质的合成等过程。试验中鉴定出与代谢有关的NADP-苹果酸酶、与蛋白质合成有关的核糖体蛋白以及具有清除活性氧功能的过氧化氢酶等基因的加强表达，这一结果与高温胁迫下沟叶结缕草的差异蛋白质组学研究的结果相一致，进一步证明了高温胁迫下沟叶结缕草通过加强代谢作用，增强蛋白质的合成作用以及提高活性氧的清除能力来提高其高温抗逆性。文中还讨论了SSH分析结果与蛋白质组学分析结果不一致的可能原因。
Temperature as a drive force of biological functions affects plant's transpiration,absorption,water potential,nearly all enzymatic reations,dormancy and growth.Thetemperature range of growth in turfgrass is narrow.The suitable temperature range of growthfor cold-season turfgrass is 15～24℃,while it is 26～32℃for warm-season turfgrass.Whenit exceeds the suitable temperature range,the natual growth and health of turfgrass will beinfluenced.Studying on the cold resistance and heat tolerance of turfgrass is a hot issue.Somestudies on the physiobiochemical mechanism of the resistance in turfgrass have been carriedout,however few information is available in the molecular mechanism of its resistance inChina.The study is to detect the physiological responses and molecular adaptationmechanism of warm-season turfgrass to temperature stress.
Two warm-type turfgrass species,the most widely planted,such as Zoysia matrella (L.)merr.and Zoysia japonica steud.,were used for the experiment to detect the physiologicalresponses to low and high temperature stress.The results showed that the contents of solublesugar,soluble protein,proline,were all increased,but the increasing changes in the contentsof soluble sugar and proline were higher in Z.matrella than in Z.japonica under the sametemperature stress.Superoxide dismutase (SOD),peroxidase (POD)and catalase (CAT),which function is clearing active oxygen,are the system of protective enzymes in cellmembrane.The study showed that the activities of SOD,POD and CAT were all builded up inthe two turfgrass species under the low and high temperature stresses.But the increased extent ofSOD and CAT in Z.matrella was much higher than that in Z.japonica,however Z.japonica showedhigher activity of POD than Z.matrella under the temperature stress.Malondialdehyde (MDA)isone major production of membrane lipid peroxidation.Its accumulation is physiologicalindicator of active oxygen toxicity.It was found that the content of MDA was insignificantlyincreased under the high temperature stress,but it was significantly enhanced under the lowtemperature stress in the two turfgrass species.The increasing extent of MDA content wasmuch higher in Z.matrella than that in Z.japonica under the low temperature stress.Photosynthetic pigments not only affect the colour and luster of turfgrass but also affect itsphotosynthesis.It was also found that low and high temperature treatment caused decreases in carotenoid (Car)、chlorophyll a (Chla)and chlorophyll b (Chlb)contents and changes inChla/Chlb.However,the two turfgrass species responded to the temperature stressdifferently.The decreasing extent of the pigments was higher in Z.japonica under thetemperature treatment.Endogenous hormones play an important role on the plant growth.Thestudy showed that the content of ABA was increased when exposed to temperature treatment,and the increased extent was much higher under 40℃treatment than under 0℃or10℃treatment in the two turfgrass species.The increased degree of ABA in Z.matrella wasmuch higher than that in Z.japonica under the same temperature treatment.It maitainedrelatively high contents of promoting hormones in the two turf grass species under the hightemperature treatment,but reverse was true in the low temperature treatment.As far as thebalance of endogenous hormones was concerned,the ratio of inhibiting hormones topromoting hormones was increased in the two turfgrass species under the temperaturetreatment.
According to the results obtained from analyzing the resistance physiological indexes,itindicated that the two turfgrass species performed stronger resistibility under the hightemperature stress than under the low temperatue stress and Z.matrella showed the strongerresistance than Z.japonica under the same temperature stress.
In this article,the warm season turfgrass,Z.matrella,which showed stronger resistancein the experiment of resistance physiology,was studied using two-dimensional gelelectrophoresis(2-DE),mass spectrometry(MS),and bioinformatics analyses to explore thedifferentially expressed proteins of the turfgrass under low temperature and high temperaturetreatments.The result indicated that there were 35 potein spots which showed differentexpression partterns after the the different treatments.Of these,26 proteins were successfullyidentified with the aid of Q-TOF MS and bioinformatics.These proteins were involved insignal transduction,metabolism,stress resistance,protein synthesis and degradation,celltransportation capacity and embryo development.There were 20 proteins identified by Q-TOFMS,of which 4 proteins were ribulose bisphosphate carboxylase large subunit,under the lowtemperature treatment.On the one hand,some of these proteins strengthened signaltransduction,active oxygen scavenging and protein defencing,which were attributed to havethe strong resistant ability in Z.matrella,on the other hand,some other proteins,which was down-regulated,the respiration was strengthen firstly and then weaken,photosythesis wasdecreased,protein degraded,and transportatio ability of cell organ reduced in Z.matrella,inturn caused the plant growth trouble.There were 19 proteins successfully identified byQ-TOF MS,of which 3 proteins were ribulose bisphosphate carboxylase large subunit,underthe high temperature treatment.These proteins strengthened signal identification,transmembrane signal transduction,photosythesis,respiration,secondary metabolism,activeoxygen scavenging,protein defencing and protein synthesis,and led to the strong resistibilityof Z.matrella under the high temperature treatment.Besides that,the relative abundance ofZLL/PNH homologous protein decreased and the decreasing extent was increased with thetime prolonged under the low temperature treatment.The similar tendency was also foundunder the high temperature stress.These suggested that it was unfavorable for the main stemmeristematic tissue development and growth either under the high temperatue stress or underthe low temperatue stress.
Further,an investigation was conducted to detect the related gene expression profiles in Z.matrella exposed to high temperature stress using suppression subtractive hybridization(SSH)method.A total of 35 ESTs from SSH-cDNA library were sequenced,and all of thesequences of inserted fragments were analyzed with the aid of bioinformatics.It obtained 26different functionally known genes.These genes were sorted into four groups such asmetabolism,gene expression regulation and signal transduction,stress resistance and proteinsynthesis.The expressions in genes of NADP-dependent malic protein,ribosomal protein S 1and catalase,which respectively involved in metabolism,protein synthesis and active oxygenscavenging,were enhanced under the high temperature stress.This result is consistent to thoseof differential proteomics research,and in turn further proved that the resistance wasstrengthened by improving metabolism,protein synthesis and the ability to scavenge activeoxygen in Z.matrella under high temperature stress.Finally,the author also discussed why alot of strengthen expression genes,which were identified in the experiment,were differentfrom outcomes of differential proteomics research in this paper.
引文
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