低夜温逆境下黄瓜果实生长与同化物代谢运转特征研究
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摘要
高效节能型日光温室是我国蔬菜栽培的主体设施,其典型的温度特征是白天温度正常,夜晚温度较低。这种温度模式对水苏糖运输型蔬菜黄瓜的果实生长和同化物运输的影响目前并不清楚。为此,我们以黄瓜耐低温品系NY-1和XC-1和不耐低温的品种津研4号为材料,设置了28℃/12℃(昼/夜)28℃/22℃(对照)两个夜温处理,研究了低夜温对黄瓜果实发育和同化物运输的影响,以期为黄瓜冬季设施专用品种的选育和冬季栽培技术的制定提供依据。
常夜温下,黄瓜果实在下午和前半夜生长较快,后半夜和上午生长较慢。低夜温胁迫下,两个耐低温品系保持相对较快果实生长速率的机制完全不同。与津研4号相比,NY-1在低夜温处理当夜果实生长较快,而XC-1的果实则在低夜温处理后的第二天白天生长较快。此时2品系同化物向果实运输的昼夜变化趋势与果实的生长速度一致。上述结果表明,在选育黄瓜耐低温品种时应注意采取多种选择策略。低夜温条件下增加果实负载可以提高第二天黄瓜源叶的光合速率,表明反馈抑制是低夜温造成黄瓜第二天光合作用下降的原因之一。
为了调查NY-1在低夜温处理时为何能保持较快的果实生长速率,观测了NY-1和津研4号低夜温下由源叶到果实路径中各组织相关糖的含量和酶的活性。结果表明,低夜温造成了2品系源叶中蔗糖、水苏糖和肌醇半乳糖苷的积累,但对果实中蔗糖、葡萄糖和果糖的含量无显著影响。黄瓜果柄是经韧皮部运输而来的水苏糖转化为蔗糖的关键部位,低夜温处理同时造成了黄瓜果柄中水苏糖含量的上升和蔗糖含量的下降,表明黄瓜果柄中由水苏糖分解为蔗糖的步骤是低夜温下黄瓜同化物向果实运输的关键障碍环节。低夜温下2品系单果株和双果株相似的果实生长速度进一步表明此时限制同化物向果实运输的关键障碍环节在库端而不是源端,同时也说明冬季黄瓜栽培时一株上应保留多个果实。进一步调查两品系果柄中由水苏糖分解为蔗糖代谢过程中各相关酶低温下的活性,未发现这些酶的耐低温能力在基因型之间存在显著差异。低夜温下NY-1果柄中ATP的含量显著高于津研4号,这可能是NY-1低夜温下能保持较快果实生长速率的重要原因。
为了调查XC-1在低夜温处理后的第二天白天保持较快果实生长速率的机制,测定了3品系成熟叶片中的各种碳水化合物的含量和关键酶的活性。结果表明,低夜温处理造成了两品系源叶中碳水化合物的积累。但在低夜温处理后的第二天上午,XC-1源叶中积累的碳水化合物主要为可运输的糖(水苏糖和蔗糖),而NY-1和津研4号则主要以淀粉(不可运输)为主。这可能是低夜温处理后的第二天白天XC-1的果实生长速度为何较NY-1和津研4号快的重要原因。
Energy-Saving Sunlight Greenhouse is very popular in China. However, the patterns of fruit growth and carbohydrate translocation of cucumber, an important stachyose-transporting vegetable under a normal–day–low–night–temperature thermoperiod (similar to the condition in Energy-Saving Sunlight Greenhouse) still remain poorly understood. In this study, One cold-sensitive cultivar (Jinyan 4) and two cold-tolerant lines (NY-1 and XC-1) of cucumber (Cucumis sativus L.) were treated with temperatures of 28°C/22°C (day/night, control) or 28°C/12°C in a 10 h photoperiod (7:00-17:00) to investigate how chilling night effect cucumber fruit growth and carbohydrate translocation. Some information helpful for cold-tolerant cucumber breeding was obtained .
Under control conditions, cucumber fruits grew fast during the afternoon and early night, and slow during the late night and morning. Under 28°C/12°C, two cold-tolerant lines adopted distinctively different mechanisms to keep relatively higher fruit growth rate. Compared to Jinyan 4, NY-1 had a higher fruit growth rate during cold nights while XC-1 fruit grew faster during the next day. Assimilate accumulation of fruits was in conformity with the growth rate under 28°C/12°C. The increase of net CO_2 assimilation rate by fruit set after a cold night indicated that feedback inhibition was partially responsible for the reduction of photosynthesis.
To investigate why NY-1 fruit grew faster than that of Jinyan 4 during cold night, carbohydrates and related enzymes were assayed from 0 h to 4 h after the start of the dark period from source leaf to fruit sink. Compared to the normal night temperature (22°C, control), sucrose, stachyose and galactinol increased in mature leaves under cold-night treatment (12°C) while sucrose, glucose and fructose in fruits remained unchanged. In peduncles, where stachyose is catabolized to sucrose after long distance transport, cold nights simultaneously induced a significant increase of stachyose (substrate) and a decrease of sucrose (product), indicating that the metabolic step from stachyose to sucrose in peduncles is crucial to translocation inhibition in cold nights. This decrease was more pronounced in the cold-sensitive cultivar. Similar growth rates of fruits on one-fruit and two-fruit plants under cold-night treatment further confirmed that it is sink activity rather than source supply limiting the source-sink translocation. No significant genotypic differences in enzyme activities involved in the stachyose-sucrose conversion, including alkalineα-galactosidase, acidα-galactosidase, galactokinase, uridine diphosphate (UDP)-galactose pyrophosphorylase, UDP–glucose-4`epimerase and sucrose synthase, were observed when assayed in an adenosine triphosphate (ATP)-rich in vitro environment. However, the ATP concentration was much higher in peduncles of the cold-tolerant line, indicating that a limiting ATP supply may be partially responsible for the stronger inhibition of the stachyose-sucrose pathway observed in the cold-sensitive cultivar (Jinyan 4).
To investigate why the fruit of XC-1 accumulate assimilate faster than that of Jinyan 4 and NY-1 in the next morning after a cold night, several carbohydrates and enzymes were assayed in mature leaves of fruit carrying nodes. Distinctly different partitioning of fixed carbon between starch and sugars (sucrose and stachyose) was observed among Jinyan 4, NY-1 and XC-1, indicating that the faster assimilate accumulation rate of XC-1 fruit in the morning after a cold night may be caused by the higher exportable carbohydrate level in the mature leaves. Higher sucrose-phosphate synthase (EC.2.4.1.14) activity constituted additional evidence that faster sucrose and stachyose biosynthesis in mature leaves was occurred in XC-1 than in Jinyan 4 and NY-1 at that time.
常夜温下,黄瓜果实在下午和前半夜生长较快,后半夜和上午生长较慢。低夜温胁迫下,两个耐低温品系保持相对较快果实生长速率的机制完全不同。与津研4号相比,NY-1在低夜温处理当夜果实生长较快,而XC-1的果实则在低夜温处理后的第二天白天生长较快。此时2品系同化物向果实运输的昼夜变化趋势与果实的生长速度一致。上述结果表明,在选育黄瓜耐低温品种时应注意采取多种选择策略。低夜温条件下增加果实负载可以提高第二天黄瓜源叶的光合速率,表明反馈抑制是低夜温造成黄瓜第二天光合作用下降的原因之一。
为了调查NY-1在低夜温处理时为何能保持较快的果实生长速率,观测了NY-1和津研4号低夜温下由源叶到果实路径中各组织相关糖的含量和酶的活性。结果表明,低夜温造成了2品系源叶中蔗糖、水苏糖和肌醇半乳糖苷的积累,但对果实中蔗糖、葡萄糖和果糖的含量无显著影响。黄瓜果柄是经韧皮部运输而来的水苏糖转化为蔗糖的关键部位,低夜温处理同时造成了黄瓜果柄中水苏糖含量的上升和蔗糖含量的下降,表明黄瓜果柄中由水苏糖分解为蔗糖的步骤是低夜温下黄瓜同化物向果实运输的关键障碍环节。低夜温下2品系单果株和双果株相似的果实生长速度进一步表明此时限制同化物向果实运输的关键障碍环节在库端而不是源端,同时也说明冬季黄瓜栽培时一株上应保留多个果实。进一步调查两品系果柄中由水苏糖分解为蔗糖代谢过程中各相关酶低温下的活性,未发现这些酶的耐低温能力在基因型之间存在显著差异。低夜温下NY-1果柄中ATP的含量显著高于津研4号,这可能是NY-1低夜温下能保持较快果实生长速率的重要原因。
为了调查XC-1在低夜温处理后的第二天白天保持较快果实生长速率的机制,测定了3品系成熟叶片中的各种碳水化合物的含量和关键酶的活性。结果表明,低夜温处理造成了两品系源叶中碳水化合物的积累。但在低夜温处理后的第二天上午,XC-1源叶中积累的碳水化合物主要为可运输的糖(水苏糖和蔗糖),而NY-1和津研4号则主要以淀粉(不可运输)为主。这可能是低夜温处理后的第二天白天XC-1的果实生长速度为何较NY-1和津研4号快的重要原因。
Energy-Saving Sunlight Greenhouse is very popular in China. However, the patterns of fruit growth and carbohydrate translocation of cucumber, an important stachyose-transporting vegetable under a normal–day–low–night–temperature thermoperiod (similar to the condition in Energy-Saving Sunlight Greenhouse) still remain poorly understood. In this study, One cold-sensitive cultivar (Jinyan 4) and two cold-tolerant lines (NY-1 and XC-1) of cucumber (Cucumis sativus L.) were treated with temperatures of 28°C/22°C (day/night, control) or 28°C/12°C in a 10 h photoperiod (7:00-17:00) to investigate how chilling night effect cucumber fruit growth and carbohydrate translocation. Some information helpful for cold-tolerant cucumber breeding was obtained .
Under control conditions, cucumber fruits grew fast during the afternoon and early night, and slow during the late night and morning. Under 28°C/12°C, two cold-tolerant lines adopted distinctively different mechanisms to keep relatively higher fruit growth rate. Compared to Jinyan 4, NY-1 had a higher fruit growth rate during cold nights while XC-1 fruit grew faster during the next day. Assimilate accumulation of fruits was in conformity with the growth rate under 28°C/12°C. The increase of net CO_2 assimilation rate by fruit set after a cold night indicated that feedback inhibition was partially responsible for the reduction of photosynthesis.
To investigate why NY-1 fruit grew faster than that of Jinyan 4 during cold night, carbohydrates and related enzymes were assayed from 0 h to 4 h after the start of the dark period from source leaf to fruit sink. Compared to the normal night temperature (22°C, control), sucrose, stachyose and galactinol increased in mature leaves under cold-night treatment (12°C) while sucrose, glucose and fructose in fruits remained unchanged. In peduncles, where stachyose is catabolized to sucrose after long distance transport, cold nights simultaneously induced a significant increase of stachyose (substrate) and a decrease of sucrose (product), indicating that the metabolic step from stachyose to sucrose in peduncles is crucial to translocation inhibition in cold nights. This decrease was more pronounced in the cold-sensitive cultivar. Similar growth rates of fruits on one-fruit and two-fruit plants under cold-night treatment further confirmed that it is sink activity rather than source supply limiting the source-sink translocation. No significant genotypic differences in enzyme activities involved in the stachyose-sucrose conversion, including alkalineα-galactosidase, acidα-galactosidase, galactokinase, uridine diphosphate (UDP)-galactose pyrophosphorylase, UDP–glucose-4`epimerase and sucrose synthase, were observed when assayed in an adenosine triphosphate (ATP)-rich in vitro environment. However, the ATP concentration was much higher in peduncles of the cold-tolerant line, indicating that a limiting ATP supply may be partially responsible for the stronger inhibition of the stachyose-sucrose pathway observed in the cold-sensitive cultivar (Jinyan 4).
To investigate why the fruit of XC-1 accumulate assimilate faster than that of Jinyan 4 and NY-1 in the next morning after a cold night, several carbohydrates and enzymes were assayed in mature leaves of fruit carrying nodes. Distinctly different partitioning of fixed carbon between starch and sugars (sucrose and stachyose) was observed among Jinyan 4, NY-1 and XC-1, indicating that the faster assimilate accumulation rate of XC-1 fruit in the morning after a cold night may be caused by the higher exportable carbohydrate level in the mature leaves. Higher sucrose-phosphate synthase (EC.2.4.1.14) activity constituted additional evidence that faster sucrose and stachyose biosynthesis in mature leaves was occurred in XC-1 than in Jinyan 4 and NY-1 at that time.
引文
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