Spd影响高温下番茄果实类胡萝卜素代谢分子机制研究及S1MAPK3的功能分析
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摘要
番茄是世界上栽培最为普遍的蔬菜作物之一。果色是衡量番茄果实品质性状的重要指标。番茄的果色主要由类胡萝卜素化合物组成,其中含量最高的是番茄红素,温度高低直接影响番茄果实颜色的深浅,夏秋季的高温常常导致番茄着色不良,延迟成熟,严重影响番茄的品质。番茄果实类胡萝卜素代谢是受多基因调控的发育过程。尽管编码类胡萝卜素合成的大多数基因都已经被分离,但目前对类胡萝卜素代谢基因的调控机制知之甚少。
多胺作为新一类植物生理活性物质,在多种逆境下也扮演着重要的角色。在植物受到环境胁迫时,作为渗透调节物质维持植物组织中阳离子和阴离子的平衡,并且它可与自由基清除剂结合通过阻止脂质氧化来抵抗氧化胁迫。本实验室先前通过转基因技术,将多胺代谢途径中SAMDC基因转入番茄,发现多胺能够提高番茄耐热性,并且果实番茄红色含量显著增加。番茄红素的增加说明多胺和番茄红素之间有一定联系,但具体关系还不太清楚,有待进一步研究。此外,高温影响番茄果实类胡萝卜素含量、多胺提高植物耐热性,以及多胺提高类胡萝卜素含量,暗示了这三者之间必然存在一定的联系。本项目拟对外源亚精胺在不同温度下影响番茄果实成熟过程、类胡萝卜素代谢生理基础以及基因表达特性进行研究；分离鉴定多胺参与调控的关键基因；研究外源多胺对抗逆性调控的生理与分子机制取得的主要结果如下：
(1)高温负调控番茄果实成熟和类胡萝卜素合成。通过外源亚精胺对三个不同温度模式下(常温26℃/20℃,亚高温33℃/27℃和高温38℃/32℃)绿熟期番茄果实进行处理,研究了在不同温度条件外源亚精胺对番茄果实成熟、果实品质以及类胡萝卜素代谢的影响。结果表明随着果实的成熟,TCI、还原糖、类胡萝卜素、可溶性固形物以及维生素含量增加,果实硬度和总酸含量降低。温度和亚精胺处理都影响了果实成熟过程和果实成熟后的物质含量。虽然亚精胺和不同温度处理对还原糖和总酸度的影响不大,但是对类胡萝卜素含量和总抗氧化活性影响很大。在常温下,外源亚精胺导致果实TCI值升高,还原糖、类胡萝卜素、抗氧化活性、乙烯产物和呼吸速率增加；可溶性固形物、总酸含量和维生素C含量降低。亚高温(33℃)处理下,还原糖、可溶性固形物、维生素C和类胡萝卜素含量降低；乙烯释放量和呼吸速率、总抗氧化活性增加。,高温和亚高温条件下外源亚精胺处理的番茄果实内抗氧化物活性提高,外源亚精胺可以缓解亚高温对番茄果实的伤害。亚精胺处理推迟了番茄果实的成熟过程。高温(38℃)处理严重地抑制了番茄成熟,从而显著地影响了番茄果实品质,但外源亚精胺并不能缓解高温对番茄果实的伤害。高温和外源亚精胺处理均能够提高内源腐胺和精胺的含量。此外,还讨论了亚精胺和高温对类胡萝卜素代谢途径的影响。
(2)为研究亚精胺调控亚高温下果实品质和类胡萝卜素代谢的作用机制,利用番茄基因芯片对常温处理、Spd处理、33℃亚高温处理以及33℃亚高温Spd共处理,四组处理下的绿熟期番茄果实转录组进行了比较分析。芯片杂交中使用10101个探针,与常温处理比较,亚高温处理的果实中127条基因表达量发生显著变化。这些变化的基因涉及到信号转导、逆境防御、氧化还原和激素代谢途径。在常温下,亚精胺处理的果实中只有34条基因表达量显著增加,涉及到初级代谢、信号转导、激素代谢、转录因子和逆境防御；55条基因表达量显著降低,涉及到能量代谢、细胞壁代谢和光合作用。功能分类和进一步的数据分析表明,在分子水平上Spd显著缓解了亚高温逆境对植物造成的伤害。
(3)为进一步研究Spd是否通过MAPK级联途径发挥调控作用,以及SlMAPK3在番茄植株抵抗亚高温逆境中的功能,我们同时构建了含组成型启动子CaMV35S和含番茄果实特异型启动子E8的过表达载体和干涉载体,利用农杆菌介导法导入番茄植株中,对获得的超量表达转基因株系进行高温、低温(4℃)、机械损伤、触摸、盐胁迫等不同逆境处理,分析SlMAPK1、SlMAPK2和SlMAPK3表达情况。发现SlMAPK1、SlMAPK2和SlMAPK3在不同的逆境处理下均呈现出表达量先升高,到达顶峰后,随之降低的趋势。与对照相比,SlMAPK1、SlMAPK2和SlMAPK3表达量显著增加,番茄SlMAPK3超量表达转基因植株抗逆性比番茄对照植株显著增加。可见SlMAPK3在番茄抗逆性中起着非常重要的作用,
Tomato (Solalnum lycopersicum L.) is an important commercial crop. Fruit colour is an important indicator of the tomato fruit quality. The major carotenoid in tomato fruits was lycopene. High temperature adversely affects the fruit quality, such as fruit colour and delayed maturity. Carotenoids formation is highly regulated by multi-genes. Most structural genes and metabolic engineering of the pathway have been characterized, but the molecular basis was unknown.
Polyamines are small ubiquitous compounds that have been implicated in the regulation of a variety of stress responses in plants. It has been suggested that PA involvement in abiotic stress adaptation could be due to their roles in osmotic adjustment, membrane stability, free-radical scavenging and regulation of stomatal movements. In our previous study, overexpression of heterologous SAMDC in tomato generally results in improving the tolerance to heat stress and carotene accumulation. But the relationship between polyamine and carotene is not sure. In addition, the carotene content affecting by high temperature, the tolerance improving by polyamine and the carotene content accumulating by polyamine, those suggest there is a positive relationship among high temperature, polyamine and carotene content. The objective of this study was to understand the respective impact of exogenous spermidine on ripening, fruit quality and carotene content with off-vine tomato under high temperature. The results are as follows:
(1) Fruit mature and carotene biosynthesis is adversely affected by high temperature. The objective of this study was to understand the respective impact of exogenous spermidine on ripening, fruit quality and carotene content with off-vine tomato under high temperature. During ripening, TCI, reducing sugars, carotenes, soluble solids content and total ascorbate increased, whereas firmness and titratable acidity decreased. Exogenous spermidine and temperature can severely affect the ripening process and the final nutrient composition of tomato fruit. Sugars and acids were not considerably modified, but carotenoids and total antioxidants were very sensitive to polyamine and temperature. Under normal temperature, exogenous spermidine enhanced fruit TCI value, reducing sugars content, total carotene content, total antioxidant activity, and ethylene production and respiration rate, while soluble solids content, titratable acidity and total ascorbic were lower than untreated fruit. Increasing temperature from26to33℃, it reduced ascorbate, reducing sugars, soluble solids and lycopene content, but enhanced ethylene production and respiration rate and total antioxidant activity. Exogenous application of spermidine could partially alleviate the effect of high temperature, spermidine-treated fruits under higher temperature exhibited higher total antioxidants than untreated fruits. High temperature (38℃) significantly delayed fruit ripening, adversely affected the fruit quality, and spermidine can not alleviate the effects. Spermidine delayed the ripening process of tomato fruits under high temperature. Meanwhile, high temperature and exogenous spermidine treatment all increased endogenous level of Put and Spm. The regulation by spermidine and temperature of the carotene biosynthesis pathways is also discussed.
(2) High temperature adversely affects quality and yield of tomato fruit. Polyamine can alleviate heat injury in plants. This study is aimed to investigate the effects of polyamine and high temperature on transcriptional profiles in ripening tomato fruit. Methods:An Affymetrix tomato microarray was used to evaluate changes in gene expresse in response to exogenous spermidine (Spd,1mmol/L) and high temperature (33℃) treatments in tomato fruits at mature green stage. Results:Of the10101tomato probe sets represented on the array,127loci were differentially expressed in high temperature-treated fruits, compared with those under normal conditions, functionally characterized by their involvement in signal transduction, defense response, oxidation reduction, and hormone responses. However, only34genes were up-regulated in Spd-treated fruits as compared with non-treated fruits, which were involved in primary metabolism, signal transduction, hormone responses, transcription factors, and stress response. Meanwhile,55genes involved in energy metabolism, cell wall metabolism, and photosynthesis were down-regulated in Spd-treated fruits. Functional categorization and the further analysis indicated that Spd might alleviate the heat stress injury during tomato fruit ripening.
(3) To study how Spd works by MAPK cascade, and the function of SIMAPK3of tomato fruits in improving tolerance to heat stress, the35S:MAPK3, E8:MAPK3,35S:mapk3-RNAi and E8: mapk3-RNAi was transform to tomato. The35S:mapk3-RNAi plants can not grow normally, while the35S.MAPK3, E8:MAPK3, and E8:mapk3-RNAi plants development normally. The plant height, leaf length, fruit weight and fruit colour were mearsured. Real-time PCR was used to determine the expression pattern of SIMAPK1, MAPK2and MAPK3in35S.MAPK3plants treated with high temperature, low temperature, touch, wounding or salt tolerance. It confirmed that the expression raised firstly, to reach the highest value, then reduced in different stress. And it was expressed at higher levels in35S:MAPK3plants than WT plants. The35S:MAPK3plants showed higher tolerance to stress than WT plants. It confirmed that SIMAPK3play important role in tomato response to different stresses.
多胺作为新一类植物生理活性物质,在多种逆境下也扮演着重要的角色。在植物受到环境胁迫时,作为渗透调节物质维持植物组织中阳离子和阴离子的平衡,并且它可与自由基清除剂结合通过阻止脂质氧化来抵抗氧化胁迫。本实验室先前通过转基因技术,将多胺代谢途径中SAMDC基因转入番茄,发现多胺能够提高番茄耐热性,并且果实番茄红色含量显著增加。番茄红素的增加说明多胺和番茄红素之间有一定联系,但具体关系还不太清楚,有待进一步研究。此外,高温影响番茄果实类胡萝卜素含量、多胺提高植物耐热性,以及多胺提高类胡萝卜素含量,暗示了这三者之间必然存在一定的联系。本项目拟对外源亚精胺在不同温度下影响番茄果实成熟过程、类胡萝卜素代谢生理基础以及基因表达特性进行研究；分离鉴定多胺参与调控的关键基因；研究外源多胺对抗逆性调控的生理与分子机制取得的主要结果如下：
(1)高温负调控番茄果实成熟和类胡萝卜素合成。通过外源亚精胺对三个不同温度模式下(常温26℃/20℃,亚高温33℃/27℃和高温38℃/32℃)绿熟期番茄果实进行处理,研究了在不同温度条件外源亚精胺对番茄果实成熟、果实品质以及类胡萝卜素代谢的影响。结果表明随着果实的成熟,TCI、还原糖、类胡萝卜素、可溶性固形物以及维生素含量增加,果实硬度和总酸含量降低。温度和亚精胺处理都影响了果实成熟过程和果实成熟后的物质含量。虽然亚精胺和不同温度处理对还原糖和总酸度的影响不大,但是对类胡萝卜素含量和总抗氧化活性影响很大。在常温下,外源亚精胺导致果实TCI值升高,还原糖、类胡萝卜素、抗氧化活性、乙烯产物和呼吸速率增加；可溶性固形物、总酸含量和维生素C含量降低。亚高温(33℃)处理下,还原糖、可溶性固形物、维生素C和类胡萝卜素含量降低；乙烯释放量和呼吸速率、总抗氧化活性增加。,高温和亚高温条件下外源亚精胺处理的番茄果实内抗氧化物活性提高,外源亚精胺可以缓解亚高温对番茄果实的伤害。亚精胺处理推迟了番茄果实的成熟过程。高温(38℃)处理严重地抑制了番茄成熟,从而显著地影响了番茄果实品质,但外源亚精胺并不能缓解高温对番茄果实的伤害。高温和外源亚精胺处理均能够提高内源腐胺和精胺的含量。此外,还讨论了亚精胺和高温对类胡萝卜素代谢途径的影响。
(2)为研究亚精胺调控亚高温下果实品质和类胡萝卜素代谢的作用机制,利用番茄基因芯片对常温处理、Spd处理、33℃亚高温处理以及33℃亚高温Spd共处理,四组处理下的绿熟期番茄果实转录组进行了比较分析。芯片杂交中使用10101个探针,与常温处理比较,亚高温处理的果实中127条基因表达量发生显著变化。这些变化的基因涉及到信号转导、逆境防御、氧化还原和激素代谢途径。在常温下,亚精胺处理的果实中只有34条基因表达量显著增加,涉及到初级代谢、信号转导、激素代谢、转录因子和逆境防御；55条基因表达量显著降低,涉及到能量代谢、细胞壁代谢和光合作用。功能分类和进一步的数据分析表明,在分子水平上Spd显著缓解了亚高温逆境对植物造成的伤害。
(3)为进一步研究Spd是否通过MAPK级联途径发挥调控作用,以及SlMAPK3在番茄植株抵抗亚高温逆境中的功能,我们同时构建了含组成型启动子CaMV35S和含番茄果实特异型启动子E8的过表达载体和干涉载体,利用农杆菌介导法导入番茄植株中,对获得的超量表达转基因株系进行高温、低温(4℃)、机械损伤、触摸、盐胁迫等不同逆境处理,分析SlMAPK1、SlMAPK2和SlMAPK3表达情况。发现SlMAPK1、SlMAPK2和SlMAPK3在不同的逆境处理下均呈现出表达量先升高,到达顶峰后,随之降低的趋势。与对照相比,SlMAPK1、SlMAPK2和SlMAPK3表达量显著增加,番茄SlMAPK3超量表达转基因植株抗逆性比番茄对照植株显著增加。可见SlMAPK3在番茄抗逆性中起着非常重要的作用,
Tomato (Solalnum lycopersicum L.) is an important commercial crop. Fruit colour is an important indicator of the tomato fruit quality. The major carotenoid in tomato fruits was lycopene. High temperature adversely affects the fruit quality, such as fruit colour and delayed maturity. Carotenoids formation is highly regulated by multi-genes. Most structural genes and metabolic engineering of the pathway have been characterized, but the molecular basis was unknown.
Polyamines are small ubiquitous compounds that have been implicated in the regulation of a variety of stress responses in plants. It has been suggested that PA involvement in abiotic stress adaptation could be due to their roles in osmotic adjustment, membrane stability, free-radical scavenging and regulation of stomatal movements. In our previous study, overexpression of heterologous SAMDC in tomato generally results in improving the tolerance to heat stress and carotene accumulation. But the relationship between polyamine and carotene is not sure. In addition, the carotene content affecting by high temperature, the tolerance improving by polyamine and the carotene content accumulating by polyamine, those suggest there is a positive relationship among high temperature, polyamine and carotene content. The objective of this study was to understand the respective impact of exogenous spermidine on ripening, fruit quality and carotene content with off-vine tomato under high temperature. The results are as follows:
(1) Fruit mature and carotene biosynthesis is adversely affected by high temperature. The objective of this study was to understand the respective impact of exogenous spermidine on ripening, fruit quality and carotene content with off-vine tomato under high temperature. During ripening, TCI, reducing sugars, carotenes, soluble solids content and total ascorbate increased, whereas firmness and titratable acidity decreased. Exogenous spermidine and temperature can severely affect the ripening process and the final nutrient composition of tomato fruit. Sugars and acids were not considerably modified, but carotenoids and total antioxidants were very sensitive to polyamine and temperature. Under normal temperature, exogenous spermidine enhanced fruit TCI value, reducing sugars content, total carotene content, total antioxidant activity, and ethylene production and respiration rate, while soluble solids content, titratable acidity and total ascorbic were lower than untreated fruit. Increasing temperature from26to33℃, it reduced ascorbate, reducing sugars, soluble solids and lycopene content, but enhanced ethylene production and respiration rate and total antioxidant activity. Exogenous application of spermidine could partially alleviate the effect of high temperature, spermidine-treated fruits under higher temperature exhibited higher total antioxidants than untreated fruits. High temperature (38℃) significantly delayed fruit ripening, adversely affected the fruit quality, and spermidine can not alleviate the effects. Spermidine delayed the ripening process of tomato fruits under high temperature. Meanwhile, high temperature and exogenous spermidine treatment all increased endogenous level of Put and Spm. The regulation by spermidine and temperature of the carotene biosynthesis pathways is also discussed.
(2) High temperature adversely affects quality and yield of tomato fruit. Polyamine can alleviate heat injury in plants. This study is aimed to investigate the effects of polyamine and high temperature on transcriptional profiles in ripening tomato fruit. Methods:An Affymetrix tomato microarray was used to evaluate changes in gene expresse in response to exogenous spermidine (Spd,1mmol/L) and high temperature (33℃) treatments in tomato fruits at mature green stage. Results:Of the10101tomato probe sets represented on the array,127loci were differentially expressed in high temperature-treated fruits, compared with those under normal conditions, functionally characterized by their involvement in signal transduction, defense response, oxidation reduction, and hormone responses. However, only34genes were up-regulated in Spd-treated fruits as compared with non-treated fruits, which were involved in primary metabolism, signal transduction, hormone responses, transcription factors, and stress response. Meanwhile,55genes involved in energy metabolism, cell wall metabolism, and photosynthesis were down-regulated in Spd-treated fruits. Functional categorization and the further analysis indicated that Spd might alleviate the heat stress injury during tomato fruit ripening.
(3) To study how Spd works by MAPK cascade, and the function of SIMAPK3of tomato fruits in improving tolerance to heat stress, the35S:MAPK3, E8:MAPK3,35S:mapk3-RNAi and E8: mapk3-RNAi was transform to tomato. The35S:mapk3-RNAi plants can not grow normally, while the35S.MAPK3, E8:MAPK3, and E8:mapk3-RNAi plants development normally. The plant height, leaf length, fruit weight and fruit colour were mearsured. Real-time PCR was used to determine the expression pattern of SIMAPK1, MAPK2and MAPK3in35S.MAPK3plants treated with high temperature, low temperature, touch, wounding or salt tolerance. It confirmed that the expression raised firstly, to reach the highest value, then reduced in different stress. And it was expressed at higher levels in35S:MAPK3plants than WT plants. The35S:MAPK3plants showed higher tolerance to stress than WT plants. It confirmed that SIMAPK3play important role in tomato response to different stresses.
引文
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