麦红吸浆虫滞育与化学物质变化研究
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摘要
滞育是昆虫中普遍存在的适应外界不良环境条件，使种群得以在复杂多变的自然界中持续繁衍和发展的一种特殊表现形式。但滞育多态现象却是麦红吸浆虫Sitodiplosis mosellana(Gehin)在长期适应自然环境的过程中形成的克服自身年发生世代数少，个体小，繁殖能力低，成虫生活力弱等不利因素，而得以在自然界中繁荣发展并偶发猖獗成灾的一个特有生存对策，也是引起麦红吸浆虫间歇性、区域性、团块分布不均匀成灾的因素之一。本文通过较全面地查阅有关文献资料，从滞育基本概念、滞育类型、滞育诱导、维持和终止、滞育与环境因素的关系、滞育昆虫的组织结构变化、滞育昆虫的生理生化特点、滞育调控机制、滞育遗传进化等8个方面对昆虫滞育研究历史和现状进行了较系统的总结和回顾；从滞育特点、滞育诱导、维持和终止、滞育形态特征、滞育生理特征、滞育化学物质变化、我国小麦吸浆虫滞育区划等6个方面对麦红吸浆虫滞育研究进行了综述。在此基础上，首次采用化学分析方法和全蛋白与同工酶电泳技术，较系统地研究了麦红吸浆虫幼虫不同时期主要化学物质和多种同工酶的变化，结果表明：
1、当年滞育幼虫与2年滞育幼虫主要化学物质含量无明显差异 对同一时期当年滞育幼虫与2年滞育幼虫总糖、甘油、蛋白质、RNA和DNA含量分析比较表明，不同滞育年限幼虫这5种物质的含量无明显差异。说明麦红吸浆虫幼虫在滞育过程中，能量消耗是非常经济的，这或许就是其个体虽然很小，但在土壤中连续滞育12年之久仍能正常化蛹羽化的重要原因之一。
2、裸露幼虫与圆茧形式不同，滞育本质相同 在田间周年系统调查的基础上，对不同时期同一时间采集的裸露幼虫与圆茧主要化学物质分析表明，同一时间的裸露幼虫与圆茧的总糖、糖元、蛋白质、RNA和DNA含量无显著差异，说明二者的滞育本质及随季节变化的特点是一致的。由此可见，裸露幼虫与圆茧虽然存在形式不同，但滞育本质是一样的。更正了以往认为裸露幼虫是解除滞育状态，圆茧是滞育状态的错误观点。
3、主要化学物质的季节变化反映了麦红吸浆虫滞育的量变特点 试验分析表明，不同时期的幼虫总糖、海藻糖、糖元、甘油、游离氨基酸、蛋白质、RNA和DNA等主要化学物质的含量及脂肪酸组分数和主要脂肪酸含量均有极显著的差异。越夏、越冬期间，总糖、海藻糖、甘油、游离氨基酸含量明显增高，秋季和春季，四者的含量明显降低；蛋白质和RNA
摘 要
含量的变化则相反，越夏、越冬期间含量降低，秋季和春季含量升高。秋季与春季比较，春
季时上述物质含量的变化幅度更大。由此可见，麦红吸浆虫滞育期间，其滞育程度随环境条
件的变化而变化，或不同个体之间滞育深浅程度有一定的差异。证明当年进入滞育的幼虫中，
部分滞育程度较浅的幼虫夏秋季化蛹羽化是完全可能的。据此结合前人室内饲养和田间调查
的结果，可将麦红吸浆虫的滞育划分为越夏滞育（over－summering diapause）、越夏至越冬滞
育（over－summer－autumn－wintering dlapause）、一次滞育（re－diapause）和延长滞育（prolonged
dlapause）4种类型。
4、部分化学物质的变化可作为衡量滞育或阶段性滞育状态的指标 麦穗幼虫和落入土中
尚未结茧的幼虫，海藻糖含量很低。结茧后，海藻糖含量明显增加，至翌年2月，一直维持
在一个较高的水平。特别是在1月和2月的结茧幼虫中，海藻糖含量最高。2月以后，海藻
糖含量又急剧下降。由此可见，海藻糖不仅与麦红吸浆虫的越冬耐寒性有一定的关系，而且
还可作为区别裸露幼虫与圆茧及指示麦红吸浆虫幼虫滞育程度的一个重要指标。
越夏、越冬幼虫苏氨酸含量高，精氨酸和 RNA含量低，RNA／DNA值小；春季解除滞育
进入活动阶段后，苏氨酸含量明显降低，甚至检测不出，精氨酸和 RNA含量明显增加，
RNA／DNA值变大。所以，苏氨酸、精氨酸、RNA含量及RNA／DNA值的大小也可作为判断
麦红吸浆虫滞育程度或滞育状态的重要指标。
眯氨酸、酪氨酸、异亮氨酸在越夏期间含量最高，明显高于其它时间的含量。这3种氨
基酸含量变化可作为麦红吸浆虫越夏滞育的指标。
谷氨酸、脯氦酸在越冬期间含量较高，特别是脯氨酸，越冬期间的含量极显著的高于越
夏期间及越冬后的发育过程中，在越冬后发育的关键时期—一3 月幼虫中尚未检测到。而亮
氨酸在越冬阶段的幼虫中含量最低。所以，这3种氨基酸含量变化可作为麦红吸浆虫越冬滞
育的指标。
越夏、越冬幼虫及越冬后未进入活动状态的圆茧脂肪酸组分中不含花生酸（C。。：。），而越
冬后进入活动状态的幼虫和麦穗幼虫脂肪酸组分中含有一定量的花生酸（C200），所以，花生
酸的有无可作为麦红吸浆虫幼虫春季解除滞育的生化指标之一。
5、不同时期幼虫全蛋白与过氧化氢酶同工酶电泳无多态性差异 分别采用PAGE和
SDS．PAGE方法对不同时期麦红吸浆虫幼虫全蛋白分析表明，PAGE分离出了9条蛋白质谱
带，SDS－PAGE分离出了 11条蛋白质谱带。但不论采用哪种方法，不同时期的幼虫谱带数完
全相同，并无本质的区别?
Diapause is a characteristic phenomenon in insects to adapt to complex and variable or adverse environmental conditions so as to reproduce and develop their populations in nature. Polymorphism of diapause is only known in population of wheat blossom midges. It is the diapause polymorphism that makes the pests overcome themselves live drawbacks such as few generations (1 generation a year generally), small size of individual, low rate of reproduction and very faint living of adult and cause serious disaster with occasional, regional and uneven dumpling characteristics. On the basis of looking up and reading lots of literatures, this paper summarized and reviewed systematically the history and current status of diapause researches in insect in eight respects as follows: elemental concept about diapause; types of diapause; induction, maintenance and termination of diapause; relationships between diapause and environment conditions; changes of organ structure in diapause insects; diapause features in physiology and biochemistry; regulation mechanism of diapause as well as hereditary evolution of diapause. The advance of diapause researches in the wheat blossom midge, Sitodiplosis mosellana (Gehin) was yet summarized and reviewed in six respects including bionomics features; mechanisms of induction, maintenance and termination; morphological characteristics; physiological features; changes of chemical substances in diapause larvae and divisions of diapause status in China. Modern analysis methods of chemical substances and polyacrylamide gum electrophoresis (PAGE) of protein and isozyme were firstly used to study changes of main chemical substances and isozymes in diapause larvae of 5". mosellana in different periods or status. The main results are as follows:
1. Obvious difference in contents of main chemical substances was not discovered in
larvae with various diapause years. Contents of total carbohydrate, glycerine, protein, RNA and DNA in larvae diapausing for 1 year or 2 years were estimated respectively. Results showed that there were not remarkable differences in content of the five chemical substances in larvae with various diapause years. It suggested that energy consumption of S. mosellana should be very economical during diapause. It might be one of important causes that the larvae of S. mosellana could pupate and emerge after 12 years of continuous diapause in soil although their size of individual is very small.
2. Non-cocooned larvae and cocooned larvae were different in appearance, but diapause essence of them was identical. The content of main chemical substances in both non-cocooned larvae and cocooned larvae was compared. The remarkable difference between non-cocooned larvae and cocooned larvae collected at the same time of various seasons was not discovered in the content of total carbohydrate, glycogen, protein, RNA and DNA. Therefore, there was of the same diapause essence and feature in both non-cocooned larvae and cocooned larvae as the seasonal change. A fault opinion that non-cocooned larvae were used to be thought as status of diapause and cocooned larvae as status of non-diapause was corrected.
3. Seasonal changes of main chemical substances displayed the quantitative features of diapause level in S. mosellana. There was remarkable difference in contents of total carbohydrate, trehalose, glycogen, glycerine, free amino acid, protein, RNA and DNA as well as main fatty acids in larvae of S. mosellana in different seasons. The contents of total carbohydrate, glycerine, free amino acid in larvae was increasing in both summer and winter and decreasing in both autumn and spring. On the contrary, the contents of protein and RNA displayed a different trend. There was more obvious change in contents of chemical substances in spring. Therefore, diapause level in larvae of S. mosellana was changed as environmental conditions and seasons changed or there were different levels of diapause in various individuals. It was proved to be possible that parts of larvae pupated and emerged
1、当年滞育幼虫与2年滞育幼虫主要化学物质含量无明显差异 对同一时期当年滞育幼虫与2年滞育幼虫总糖、甘油、蛋白质、RNA和DNA含量分析比较表明，不同滞育年限幼虫这5种物质的含量无明显差异。说明麦红吸浆虫幼虫在滞育过程中，能量消耗是非常经济的，这或许就是其个体虽然很小，但在土壤中连续滞育12年之久仍能正常化蛹羽化的重要原因之一。
2、裸露幼虫与圆茧形式不同，滞育本质相同 在田间周年系统调查的基础上，对不同时期同一时间采集的裸露幼虫与圆茧主要化学物质分析表明，同一时间的裸露幼虫与圆茧的总糖、糖元、蛋白质、RNA和DNA含量无显著差异，说明二者的滞育本质及随季节变化的特点是一致的。由此可见，裸露幼虫与圆茧虽然存在形式不同，但滞育本质是一样的。更正了以往认为裸露幼虫是解除滞育状态，圆茧是滞育状态的错误观点。
3、主要化学物质的季节变化反映了麦红吸浆虫滞育的量变特点 试验分析表明，不同时期的幼虫总糖、海藻糖、糖元、甘油、游离氨基酸、蛋白质、RNA和DNA等主要化学物质的含量及脂肪酸组分数和主要脂肪酸含量均有极显著的差异。越夏、越冬期间，总糖、海藻糖、甘油、游离氨基酸含量明显增高，秋季和春季，四者的含量明显降低；蛋白质和RNA
摘 要
含量的变化则相反，越夏、越冬期间含量降低，秋季和春季含量升高。秋季与春季比较，春
季时上述物质含量的变化幅度更大。由此可见，麦红吸浆虫滞育期间，其滞育程度随环境条
件的变化而变化，或不同个体之间滞育深浅程度有一定的差异。证明当年进入滞育的幼虫中，
部分滞育程度较浅的幼虫夏秋季化蛹羽化是完全可能的。据此结合前人室内饲养和田间调查
的结果，可将麦红吸浆虫的滞育划分为越夏滞育（over－summering diapause）、越夏至越冬滞
育（over－summer－autumn－wintering dlapause）、一次滞育（re－diapause）和延长滞育（prolonged
dlapause）4种类型。
4、部分化学物质的变化可作为衡量滞育或阶段性滞育状态的指标 麦穗幼虫和落入土中
尚未结茧的幼虫，海藻糖含量很低。结茧后，海藻糖含量明显增加，至翌年2月，一直维持
在一个较高的水平。特别是在1月和2月的结茧幼虫中，海藻糖含量最高。2月以后，海藻
糖含量又急剧下降。由此可见，海藻糖不仅与麦红吸浆虫的越冬耐寒性有一定的关系，而且
还可作为区别裸露幼虫与圆茧及指示麦红吸浆虫幼虫滞育程度的一个重要指标。
越夏、越冬幼虫苏氨酸含量高，精氨酸和 RNA含量低，RNA／DNA值小；春季解除滞育
进入活动阶段后，苏氨酸含量明显降低，甚至检测不出，精氨酸和 RNA含量明显增加，
RNA／DNA值变大。所以，苏氨酸、精氨酸、RNA含量及RNA／DNA值的大小也可作为判断
麦红吸浆虫滞育程度或滞育状态的重要指标。
眯氨酸、酪氨酸、异亮氨酸在越夏期间含量最高，明显高于其它时间的含量。这3种氨
基酸含量变化可作为麦红吸浆虫越夏滞育的指标。
谷氨酸、脯氦酸在越冬期间含量较高，特别是脯氨酸，越冬期间的含量极显著的高于越
夏期间及越冬后的发育过程中，在越冬后发育的关键时期—一3 月幼虫中尚未检测到。而亮
氨酸在越冬阶段的幼虫中含量最低。所以，这3种氨基酸含量变化可作为麦红吸浆虫越冬滞
育的指标。
越夏、越冬幼虫及越冬后未进入活动状态的圆茧脂肪酸组分中不含花生酸（C。。：。），而越
冬后进入活动状态的幼虫和麦穗幼虫脂肪酸组分中含有一定量的花生酸（C200），所以，花生
酸的有无可作为麦红吸浆虫幼虫春季解除滞育的生化指标之一。
5、不同时期幼虫全蛋白与过氧化氢酶同工酶电泳无多态性差异 分别采用PAGE和
SDS．PAGE方法对不同时期麦红吸浆虫幼虫全蛋白分析表明，PAGE分离出了9条蛋白质谱
带，SDS－PAGE分离出了 11条蛋白质谱带。但不论采用哪种方法，不同时期的幼虫谱带数完
全相同，并无本质的区别?
Diapause is a characteristic phenomenon in insects to adapt to complex and variable or adverse environmental conditions so as to reproduce and develop their populations in nature. Polymorphism of diapause is only known in population of wheat blossom midges. It is the diapause polymorphism that makes the pests overcome themselves live drawbacks such as few generations (1 generation a year generally), small size of individual, low rate of reproduction and very faint living of adult and cause serious disaster with occasional, regional and uneven dumpling characteristics. On the basis of looking up and reading lots of literatures, this paper summarized and reviewed systematically the history and current status of diapause researches in insect in eight respects as follows: elemental concept about diapause; types of diapause; induction, maintenance and termination of diapause; relationships between diapause and environment conditions; changes of organ structure in diapause insects; diapause features in physiology and biochemistry; regulation mechanism of diapause as well as hereditary evolution of diapause. The advance of diapause researches in the wheat blossom midge, Sitodiplosis mosellana (Gehin) was yet summarized and reviewed in six respects including bionomics features; mechanisms of induction, maintenance and termination; morphological characteristics; physiological features; changes of chemical substances in diapause larvae and divisions of diapause status in China. Modern analysis methods of chemical substances and polyacrylamide gum electrophoresis (PAGE) of protein and isozyme were firstly used to study changes of main chemical substances and isozymes in diapause larvae of 5". mosellana in different periods or status. The main results are as follows:
1. Obvious difference in contents of main chemical substances was not discovered in
larvae with various diapause years. Contents of total carbohydrate, glycerine, protein, RNA and DNA in larvae diapausing for 1 year or 2 years were estimated respectively. Results showed that there were not remarkable differences in content of the five chemical substances in larvae with various diapause years. It suggested that energy consumption of S. mosellana should be very economical during diapause. It might be one of important causes that the larvae of S. mosellana could pupate and emerge after 12 years of continuous diapause in soil although their size of individual is very small.
2. Non-cocooned larvae and cocooned larvae were different in appearance, but diapause essence of them was identical. The content of main chemical substances in both non-cocooned larvae and cocooned larvae was compared. The remarkable difference between non-cocooned larvae and cocooned larvae collected at the same time of various seasons was not discovered in the content of total carbohydrate, glycogen, protein, RNA and DNA. Therefore, there was of the same diapause essence and feature in both non-cocooned larvae and cocooned larvae as the seasonal change. A fault opinion that non-cocooned larvae were used to be thought as status of diapause and cocooned larvae as status of non-diapause was corrected.
3. Seasonal changes of main chemical substances displayed the quantitative features of diapause level in S. mosellana. There was remarkable difference in contents of total carbohydrate, trehalose, glycogen, glycerine, free amino acid, protein, RNA and DNA as well as main fatty acids in larvae of S. mosellana in different seasons. The contents of total carbohydrate, glycerine, free amino acid in larvae was increasing in both summer and winter and decreasing in both autumn and spring. On the contrary, the contents of protein and RNA displayed a different trend. There was more obvious change in contents of chemical substances in spring. Therefore, diapause level in larvae of S. mosellana was changed as environmental conditions and seasons changed or there were different levels of diapause in various individuals. It was proved to be possible that parts of larvae pupated and emerged
引文
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