CO_2升高和短期高温胁迫对玉米幼苗生理生化指标的影响
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摘要
为了深入系统地研究C02浓度升高条件下,短暂高温对玉米苗期的光合生理生物化学影响,本课题在不同章节按以下思路展开：(1)确定引起光合作用呈现可逆性恢复而不影响叶片形态变化的高温阈值。同时确定高温胁迫处理时间；探讨生长期间的不同CO2浓度处理对玉米幼苗植物学性状和光合生理学的影响,以期了解玉米的哪些性状会对C02升高出现环境适应性反应；(2)提出假设,高温胁迫伴随饱和水汽压差的增加导致叶片蒸腾速率和气孔导度的降低,过量的蒸发降低水分利用效率可导致叶片干燥,进而C02有利于降低蒸腾和气孔导度,进而保持较高的叶片水分状态。(3)为了进一步分析C02对光合耐温性的影响,试验设置在室内和田间条件下,重点研究CO2升高是否会对高温处理的不同耐温性玉米品种光合生理和水分生理具有调节作用；高温下,光合速率的降低是否与光合酶活性、基因转录水平、细胞膜热稳定性和热激蛋白表达有关。(4)进一步分析细胞代谢物组分对高温胁迫的响应及CO2参与的条件作用。根据以上思路和研究方法,本课题主要的结论如下：
(1)45℃高温是引起光合速率降低,并在24h期间出现明显恢复的最佳温度。结果也表明45℃高温胁迫15分钟会立即导致光合速率的降低,并在胁迫2小时后光合速率趋于平稳。本研究故选取2小时作为温度胁迫时间用于与CO2的互作研究。CO2升高对于根、茎和叶干重均无明显影响。但C02升高会增加玉米幼苗的叶面积,这在B76玉米品种中表现更为明显(升高28.8%)。SQ具有较快的生长速度。田间条件下,CO2升高有利于3个玉米品种的NADP-MDH酶活性升高。C02升高使B76和SQ的NADP-MDH酶活性分别升高了20%和31.6%。而CO2浓度升高会降低PEPCase的活性。
(2)CO2升高不会缓解热胁迫期间的叶片水分潜势来改善光合。热胁迫期间的不同CO2浓度对光合耐热性有显著影响,生长期间不同的CO2浓度处理对光合热耐性无显著影响。热胁迫处理消除CO2效应。当叶片温度恢复到35℃时,气孔导度和胞间CO2浓度相似或更高最初值。
(3)B76和B106在田间和室内条件下表现出相反的光合热耐性。室内条件下,B76表现出高度的耐热性。但在田间条件表现较差的光合热耐性。B106则相反。蒸腾速率和气孔导度不同生长环境下也表现相反的变化趋势。田间条件下,高温胁迫降低了叶片蒸腾和气孔导度。室内条件则相反。C02升高降低了B76的蒸腾速率和气孔导度,改善了叶片水分潜势,同时一定程度上改善了叶片光合热耐性。热胁迫处理诱导的B106叶片相对伤害均高于B76。表明B106具有较低细胞膜热稳定性。PEPC在酶活性和基因转录丰度方面均具有更好的高温耐性。热胁迫导致了所有热激蛋白基因表达的上调。而各个热激蛋白无C02效应。热胁迫所诱导的热激蛋白70表达丰度在SQ中达到250倍,比在同等条件下的B106高出66.7%。B76中,Hsp101和Hsp82的恢复动态与光合动态具有高度线性相关。
(4)在热胁迫下,总非结构性碳水化合物含量在所有玉米品种中降低。根据代谢分析结果表明C02富集增加了B76中25个代谢物成分。在2个C02处理下,丙酮酸和苹果酸受到热胁迫的最大影响,说明B76在热胁迫期间光合的降低可能与三羧酸循环进程被高温扰断有关。热胁迫诱导亮氨酸、甘氨酸、丝氨酸、脯氨酸、谷氨酸含量、葡萄糖和果糖的升高。热胁迫同时诱导了异亮氨酸、缬氨酸、天冬氨酸含量和淀粉的降低。
To further systematically explore the photosynthetic physiology and biochemistry in response to short-term high temperature under elevated CO2in maize seedlings, this dissertation based on different chapters in turn were stretched out as following text,(1) Examine the high temperature threshold that can induce complete reverse of photosynthesis maintaining intact leave phenotypes. Focus on botanical characteristic and photosynthetic variables in response to different CO2levels for maize seedlings during growth period, to understand which traits exhibited acclimation to elevated CO2.(2) A hypothesis to prove stomatal conduct ances and transpiration increased with vapour pressure deficit and high temperature stress could be mitigated via elevated CO2effects on pore patern and water status, plants were subjected to combination of heat stress treatments and four CO2conditions (3) To further analysis on effects of CO2on phototysnthetic thermotolerance, plants were grown at indoor chambers and field. This study tested weather elevated CO2have modification on different maize cultivars that have contrasting thermotolerance in terms of photosynthetic variable and water status. Based on primary technical system and theory base, the study analyzed vitro activities of C4key photosynthetic enzymes, their transcriptional abundance, and integrity of cellular membrane.(4) to explore stressed-protein involved disruption and repairing by abiotic stress. Further test cellular metabolic reprogramming in response to heat stress and CO2effects was done. Based technical base and methods, this dissertation concluded as below:
(1) High temperature45℃induced the decrease of photosynthesis, and was the best candidate temperature for abiostic temperature for obvious recovery after heat stress24h. There was no difference between2h and4h regarding the decrease of photosynthesis. This study investigated the interactive effects of heat treated time and CO2. Elevated CO2has no significance effects on root, stem and leaves, but enhanced leaf areas of maize seedlings, especially for B76(28.8%). SQ has fast growth developments. Activities of NADP-MDH increased under elevated CO2. Activities of NADP-MDH increased20%and31.6%for B76and SQ under elevated CO2. The activities of PEPCase decreased under elevated CO2.
(2) Elevated CO2cannot mitigate the water potential during heat stress to protect the damage of photosynthesis. Effects of different CO2on photosynthetic thermotolerance have no significance difference. Heat treatment eliminated CO2effects. When leaf temperature recovered to35℃, gs and Ci maintained similar or higher values than initial values.
(3) An opposite performances of photosynthetic tolerance was observed between field and indoor conditions. While under indoor, B76has more tolerance to heat stress. But under field conditions, photosynthetic tolerance was decreasing rather than B106. Transpiration and stomatal conductances performed opposite trends. Under field conditions, high temperature decreased transpiration and stomatal condutances rather than indoor conditions. Elevated CO2decreased the transpiration and stomatal conductances of B76, improving water potential of leaves as well as tolerance to heat stress. Heat stress induced relative injuries of B106were higher than B76suggesting B106has lower thermostability of cellular membrane. Activates of PEPCase and its gene expression were upregulated. But each heat shock protein has no CO2effects. Heat stress induced250times upregulation of HSP70expression in SQ,66.7%higher than B106at same conditions. In B76, the patterns of Hsp101and Hsp82have correlation with photosynthesis.
(4) Under heat stress, total non-structural carbonhydrates decreased in all of maize cultivars. According to metabolic analysis, CO2enrichments increased25metabolites in B76. Under two CO2treatments, pyruvic acids and malic acids were influenced by heat treatments, suggesting the decrease of photosynthesis in B76probably was attributed to disruption of TCA cycles. Heat stress induced lectine, glycine, serine, proline, glutamate, fructose and glucose as well as decrease of isolectine, valine, aspirate and starch.
(1)45℃高温是引起光合速率降低,并在24h期间出现明显恢复的最佳温度。结果也表明45℃高温胁迫15分钟会立即导致光合速率的降低,并在胁迫2小时后光合速率趋于平稳。本研究故选取2小时作为温度胁迫时间用于与CO2的互作研究。CO2升高对于根、茎和叶干重均无明显影响。但C02升高会增加玉米幼苗的叶面积,这在B76玉米品种中表现更为明显(升高28.8%)。SQ具有较快的生长速度。田间条件下,CO2升高有利于3个玉米品种的NADP-MDH酶活性升高。C02升高使B76和SQ的NADP-MDH酶活性分别升高了20%和31.6%。而CO2浓度升高会降低PEPCase的活性。
(2)CO2升高不会缓解热胁迫期间的叶片水分潜势来改善光合。热胁迫期间的不同CO2浓度对光合耐热性有显著影响,生长期间不同的CO2浓度处理对光合热耐性无显著影响。热胁迫处理消除CO2效应。当叶片温度恢复到35℃时,气孔导度和胞间CO2浓度相似或更高最初值。
(3)B76和B106在田间和室内条件下表现出相反的光合热耐性。室内条件下,B76表现出高度的耐热性。但在田间条件表现较差的光合热耐性。B106则相反。蒸腾速率和气孔导度不同生长环境下也表现相反的变化趋势。田间条件下,高温胁迫降低了叶片蒸腾和气孔导度。室内条件则相反。C02升高降低了B76的蒸腾速率和气孔导度,改善了叶片水分潜势,同时一定程度上改善了叶片光合热耐性。热胁迫处理诱导的B106叶片相对伤害均高于B76。表明B106具有较低细胞膜热稳定性。PEPC在酶活性和基因转录丰度方面均具有更好的高温耐性。热胁迫导致了所有热激蛋白基因表达的上调。而各个热激蛋白无C02效应。热胁迫所诱导的热激蛋白70表达丰度在SQ中达到250倍,比在同等条件下的B106高出66.7%。B76中,Hsp101和Hsp82的恢复动态与光合动态具有高度线性相关。
(4)在热胁迫下,总非结构性碳水化合物含量在所有玉米品种中降低。根据代谢分析结果表明C02富集增加了B76中25个代谢物成分。在2个C02处理下,丙酮酸和苹果酸受到热胁迫的最大影响,说明B76在热胁迫期间光合的降低可能与三羧酸循环进程被高温扰断有关。热胁迫诱导亮氨酸、甘氨酸、丝氨酸、脯氨酸、谷氨酸含量、葡萄糖和果糖的升高。热胁迫同时诱导了异亮氨酸、缬氨酸、天冬氨酸含量和淀粉的降低。
To further systematically explore the photosynthetic physiology and biochemistry in response to short-term high temperature under elevated CO2in maize seedlings, this dissertation based on different chapters in turn were stretched out as following text,(1) Examine the high temperature threshold that can induce complete reverse of photosynthesis maintaining intact leave phenotypes. Focus on botanical characteristic and photosynthetic variables in response to different CO2levels for maize seedlings during growth period, to understand which traits exhibited acclimation to elevated CO2.(2) A hypothesis to prove stomatal conduct ances and transpiration increased with vapour pressure deficit and high temperature stress could be mitigated via elevated CO2effects on pore patern and water status, plants were subjected to combination of heat stress treatments and four CO2conditions (3) To further analysis on effects of CO2on phototysnthetic thermotolerance, plants were grown at indoor chambers and field. This study tested weather elevated CO2have modification on different maize cultivars that have contrasting thermotolerance in terms of photosynthetic variable and water status. Based on primary technical system and theory base, the study analyzed vitro activities of C4key photosynthetic enzymes, their transcriptional abundance, and integrity of cellular membrane.(4) to explore stressed-protein involved disruption and repairing by abiotic stress. Further test cellular metabolic reprogramming in response to heat stress and CO2effects was done. Based technical base and methods, this dissertation concluded as below:
(1) High temperature45℃induced the decrease of photosynthesis, and was the best candidate temperature for abiostic temperature for obvious recovery after heat stress24h. There was no difference between2h and4h regarding the decrease of photosynthesis. This study investigated the interactive effects of heat treated time and CO2. Elevated CO2has no significance effects on root, stem and leaves, but enhanced leaf areas of maize seedlings, especially for B76(28.8%). SQ has fast growth developments. Activities of NADP-MDH increased under elevated CO2. Activities of NADP-MDH increased20%and31.6%for B76and SQ under elevated CO2. The activities of PEPCase decreased under elevated CO2.
(2) Elevated CO2cannot mitigate the water potential during heat stress to protect the damage of photosynthesis. Effects of different CO2on photosynthetic thermotolerance have no significance difference. Heat treatment eliminated CO2effects. When leaf temperature recovered to35℃, gs and Ci maintained similar or higher values than initial values.
(3) An opposite performances of photosynthetic tolerance was observed between field and indoor conditions. While under indoor, B76has more tolerance to heat stress. But under field conditions, photosynthetic tolerance was decreasing rather than B106. Transpiration and stomatal conductances performed opposite trends. Under field conditions, high temperature decreased transpiration and stomatal condutances rather than indoor conditions. Elevated CO2decreased the transpiration and stomatal conductances of B76, improving water potential of leaves as well as tolerance to heat stress. Heat stress induced relative injuries of B106were higher than B76suggesting B106has lower thermostability of cellular membrane. Activates of PEPCase and its gene expression were upregulated. But each heat shock protein has no CO2effects. Heat stress induced250times upregulation of HSP70expression in SQ,66.7%higher than B106at same conditions. In B76, the patterns of Hsp101and Hsp82have correlation with photosynthesis.
(4) Under heat stress, total non-structural carbonhydrates decreased in all of maize cultivars. According to metabolic analysis, CO2enrichments increased25metabolites in B76. Under two CO2treatments, pyruvic acids and malic acids were influenced by heat treatments, suggesting the decrease of photosynthesis in B76probably was attributed to disruption of TCA cycles. Heat stress induced lectine, glycine, serine, proline, glutamate, fructose and glucose as well as decrease of isolectine, valine, aspirate and starch.
引文
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