棉蚜（Aphis gossypii）对温度和光周期适应对策的进化
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摘要
棉蚜(Aphis gossypii Glover)属同翅目(Homoptera)蚜科(Aphididae)蚜属(Aphis)，能寄生74科285种植物，是一种世界性的害虫。棉蚜是我国棉花、瓜类等作物的重要害虫，其在我国多个省份均能进行危害，每年对我国的农业生产造成重要损失。进入二十世纪80年代，棉蚜更已成为我国华北及西北半干旱地区的主要害虫。1986年吐鲁番棉蚜大暴发，直接经济损失达2000万元；1988年新疆阿克苏地区棉蚜大发生，面积达450000亩，经济损失严重；1994年新疆北疆棉区棉蚜大暴发，对棉花生产造成严重损失。预计2003年棉蚜在全国的发生面积将高达6200多万亩次。
棉蚜的大发生与农药的不合理使用有着一定的关系，同时棉蚜种群较强的异质性，较大的遗传变异度以及对生态系统极强的适应能力，也都为其广泛分布、危害成灾提供了便利条件。世界性分布的害虫，其地理分布区域可能跨越不同的气候带，而不同的气候带中又存在着各异的气候条件，因而生物在生存过程中，首先要克服的便是环境问题。棉蚜在世界各地都有分布，这说明其对周围环境具有很强的适应能力，同时也可能已进化出了一套相对完善的生态适应对策，而这种生态适应对策正可能是导致其广泛成灾的内在要素，因此展开相关方面的研究显得尤为重要。本次实验通过比较研究的方法，对越南种群(YN)、海南种群(HN)、北京种群(BJ)和新疆种群(XJ)四个地理种群棉蚜的形态、生长发育参数作了系统研究；同时利用低温、短日照条件对各地理种群棉蚜进行了性蚜的诱导，主要实验结果如下：
1．光周期L：D=16h：8h，相对湿度70％条件下，设置35℃、30℃、25℃、20℃、15℃五个温度梯度，研究了四个地理种群棉蚜体型和体色的变化。结果表明，不同气候带的棉蚜其在体型和体色特征方面存在着分化。①海南棉蚜种群与北京种群和新疆种群在头宽、前足腿节长、前足胫节长、中足腿节长、中足胫节长、后足腿节长、后足胫节长和腹管长方面均存在显著差异。②棉蚜成蚜和若蚜体型有随温度降低逐渐增大的趋势。在低温15℃，越南、海南和北京种群棉蚜成蚜体表面积有随取样点纬度的升高而明显增大的趋势。③在试验温度下，各种群棉蚜由若蚜发育至成蚜过程中，绿色蚜所占比例均有所上升，即随着生长发育的进行，棉蚜体色在世代内有逐渐加深的趋势。20℃和15℃，各种群初孵若蚜中绿色蚜所占比例为0，而当其发育至成蚜后则均出现一定比例的绿色蚜。在20℃，北京种群棉蚜发育至成蚜后绿色蚜比例上升为68.89％，15℃上升为87.50％，均高于其它种群。即表现为来自高纬度地区的棉蚜种群，在低温的条件下，其体色加深的速率相对较快。
2．光周期L：D=16h：8h，相对湿度70％条件下，设置35℃、30℃、25℃、20℃、15℃五个温度梯度，应用实验种群生命表技术，比较了不同气候带棉蚜(越南种群、北京种群、新疆种群)的存活、生长发育、繁殖及内禀增长力参数的差异。结果表明：①棉蚜在15℃-30℃可正常存活，温度高达35℃则不能正常存活；②棉蚜幼虫历期种群间差异不显著，种群内有随温度降低而增长的趋势；成虫历期30℃越南种群最长，15℃新疆种群＞北京种群＞越南种群；③产蚜量30℃越南种群最大，15℃新疆种群＞北京种群＞越南种群。即15℃新疆种群、北京种群、越南种群之间，棉蚜的成虫历期和产
东北农业大学农学硕士学位论文
蚜量均有随种群取样点纬度的增加而增大的趋势；
3．实验室中，18’C，L：D＝8：16和L：D＝10：14条件下，对越南种群（YN）、海南
种群（HN）、北京种群（BJ）、新疆种群（XJ）四个地理种群的棉蚜进行了性蚜的诱导。
结果表明：①经人工诱导，YN、删没有性蚜出现，而BJ、XJ出现性蚜并成功产下受精
卵：②棉蚜人工诱导的总性蚜率有随产仔天数的增长而增加的趋势。
根据以上试验结果，同时结合相关报道，可得出以下结论：棉蚜在对所处环境的长
期适应过程中，为了生存和种群的延续，也己相应地进化出了一系列的生态适应对策。
在形态学方面，可通过采用调节体表面积和体色的对策来适应周围环境条件的变化，低
纬度地区棉蚜，一般表面积较小、体色较淡；而高纬度地区棉蚜为了有效地获得生存所
需的积温，一般体表面积较大、体色较深。同时在对环境的长期适应过程中，棉蚜成虫
的寿命和产卵量也呈现一定程度的分化。低纬度地区地处热带常年高温，这使得生活在
其中的棉蚜种群经过长期的进化作用，对高温己产生了一定的适应性，因而在高温的条
件下，成虫的寿命较长，产卵量都较大；而高纬度地区的棉蚜，由于对低温的适应，反
而是在低温的条件下，呈现相应的变化。在低纬度地区，由于环境条件适宜，棉蚜终生
行孤雌生殖，不出现有性世代；而在高纬度地区，为了度过寒冷的冬季，棉蚜主要以有
性世代产生的受精卵越冬。可以看出，棉蚜对于不同的生态环境都有其独特的适应对策，
而所有这些生态适应对策，都为其广泛分布，发生危害提供了便利条件。
Cotton aphid Aphis gossypii (Homoptera: Aphididae), an important agricultural pest, its distribution extends from 60 N to 40 S in the world. This insect is capable to survive under different climatic conditions due to its wider adaptability. In summer, this insect reproduces parthenogenetically from generation to generation, after the late autumn it produces sexual aphids to lay eggs to over-winter. This paper presents studies on ecological adaptability of Aphis gossypii under different temperature and photoperiod ranges in the laboratory.
Adults of Aphis gossypii were collected from cotton plants in Ho Chi Minh city of Vietnam (106.42 E, 10.45 N), Beijing (116.5 E, 39.9 N), Shihezi city of Xinjiang Municipality (85 E, 45 N) and Sanya city of Hainan Province (109.5 E, 18.2 N) in 2001 and these adult populations were named as YN, BJ, XJ and HN respectively. Each aphid culture was maintained on cotton seedlings at 22 C, 70%RH, and a photoperiod of 14:10 (L:D). The morphological variation of growth and development and relationship between emergence of sexual morphs and photoperiod (L:D=8:16 and L:D=10:14) at 18 C as well as the life parameters of these populations at different temperatures were evaluated.
In the 1st part of the study with asexual aphids, fifty aphids from each of the 4 populations (i.e. YN, HN, BJ and XJ) were placed at various temperatures (15, 20, 25, 30 and 35 C respectively) maintaining 70%RH, and photoperiod L:D=16:8. Each aphid from respective population was fed on cotton cotyledon in 25ml glass beaker with 10ml-dissolved medium at the bottom. The newly produced aphids were transferred on cotyledon, and the cotton cotyledon was put on the surface of culture medium tightly. The newly produced nymphs were picked out everyday, but the adults were allowed to remain in the same container with replacement of fresh cotyledon and medium as and when required (Leaf Wafer Method). In the experiment, forty newly produced nymphs and the dead adults were all kept. Morphological parameters, such as foreleg femur length, foreleg tibia length, midleg femur length, midleg tibia length, hindleg femur length, hindleg tibia length, body length and body width were studied.
In the 2nd part of the study with sexual aphids (collected from cotton plants), forty 4th instar aphids (G0) were transferred into environment chamber and allowed to breed using LWM. After 24 hours, the G0 adult aphids were removed, and forty new nymphs (G1) from G0 generation were introduced to breed until generation three (G3). The G2 nymphs were kept on feeding, which were produced at the 2nd, 7th, 12th and 17th day separately. During the growth period, the number of apteral and alate aphids was checked and the alate ones were transferred to cotton plants having 3 to 4 leaves to get G3 generation. The cotton plants were enclosed with transparent plastic sleeves, which were covered at the top with gauze. The rates
of development of three types of aphids (1) virginoparae cotton aphid in G3 with reproductive potential, (2) male alate aphid with genetalia without reproductive potential and, (3) female apteral aphid having eggs ] were observed.
In the 3rd part of the study, fifty aphids from each of the 3 populations (i.e. YN, BJ and XJ) were placed at various temperatures (15, 20, 25, 30 and 35 C respectively) maintaining 70%RH, and photoperiod L:D=16:8. Each aphid from respective population was breed with LWM. Ecdysis, reproduction and death of aphid were observed everyday until all aphid adults died.
The results showed that there were significant differences of morphology and color while they developed under various temperatures. Length of foreleg femur, foreleg tibia, midleg femur, midleg tibia, hindleg femur and hindleg tibia, head width and cornicles length of aphid from HN showed significant differences with those from BJ and XJ population. The surface area of both adult and larval aphids gradually increased with decreased temperature. It was found that there were also observed significant differences in surface area among Y
棉蚜的大发生与农药的不合理使用有着一定的关系，同时棉蚜种群较强的异质性，较大的遗传变异度以及对生态系统极强的适应能力，也都为其广泛分布、危害成灾提供了便利条件。世界性分布的害虫，其地理分布区域可能跨越不同的气候带，而不同的气候带中又存在着各异的气候条件，因而生物在生存过程中，首先要克服的便是环境问题。棉蚜在世界各地都有分布，这说明其对周围环境具有很强的适应能力，同时也可能已进化出了一套相对完善的生态适应对策，而这种生态适应对策正可能是导致其广泛成灾的内在要素，因此展开相关方面的研究显得尤为重要。本次实验通过比较研究的方法，对越南种群(YN)、海南种群(HN)、北京种群(BJ)和新疆种群(XJ)四个地理种群棉蚜的形态、生长发育参数作了系统研究；同时利用低温、短日照条件对各地理种群棉蚜进行了性蚜的诱导，主要实验结果如下：
1．光周期L：D=16h：8h，相对湿度70％条件下，设置35℃、30℃、25℃、20℃、15℃五个温度梯度，研究了四个地理种群棉蚜体型和体色的变化。结果表明，不同气候带的棉蚜其在体型和体色特征方面存在着分化。①海南棉蚜种群与北京种群和新疆种群在头宽、前足腿节长、前足胫节长、中足腿节长、中足胫节长、后足腿节长、后足胫节长和腹管长方面均存在显著差异。②棉蚜成蚜和若蚜体型有随温度降低逐渐增大的趋势。在低温15℃，越南、海南和北京种群棉蚜成蚜体表面积有随取样点纬度的升高而明显增大的趋势。③在试验温度下，各种群棉蚜由若蚜发育至成蚜过程中，绿色蚜所占比例均有所上升，即随着生长发育的进行，棉蚜体色在世代内有逐渐加深的趋势。20℃和15℃，各种群初孵若蚜中绿色蚜所占比例为0，而当其发育至成蚜后则均出现一定比例的绿色蚜。在20℃，北京种群棉蚜发育至成蚜后绿色蚜比例上升为68.89％，15℃上升为87.50％，均高于其它种群。即表现为来自高纬度地区的棉蚜种群，在低温的条件下，其体色加深的速率相对较快。
2．光周期L：D=16h：8h，相对湿度70％条件下，设置35℃、30℃、25℃、20℃、15℃五个温度梯度，应用实验种群生命表技术，比较了不同气候带棉蚜(越南种群、北京种群、新疆种群)的存活、生长发育、繁殖及内禀增长力参数的差异。结果表明：①棉蚜在15℃-30℃可正常存活，温度高达35℃则不能正常存活；②棉蚜幼虫历期种群间差异不显著，种群内有随温度降低而增长的趋势；成虫历期30℃越南种群最长，15℃新疆种群＞北京种群＞越南种群；③产蚜量30℃越南种群最大，15℃新疆种群＞北京种群＞越南种群。即15℃新疆种群、北京种群、越南种群之间，棉蚜的成虫历期和产
东北农业大学农学硕士学位论文
蚜量均有随种群取样点纬度的增加而增大的趋势；
3．实验室中，18’C，L：D＝8：16和L：D＝10：14条件下，对越南种群（YN）、海南
种群（HN）、北京种群（BJ）、新疆种群（XJ）四个地理种群的棉蚜进行了性蚜的诱导。
结果表明：①经人工诱导，YN、删没有性蚜出现，而BJ、XJ出现性蚜并成功产下受精
卵：②棉蚜人工诱导的总性蚜率有随产仔天数的增长而增加的趋势。
根据以上试验结果，同时结合相关报道，可得出以下结论：棉蚜在对所处环境的长
期适应过程中，为了生存和种群的延续，也己相应地进化出了一系列的生态适应对策。
在形态学方面，可通过采用调节体表面积和体色的对策来适应周围环境条件的变化，低
纬度地区棉蚜，一般表面积较小、体色较淡；而高纬度地区棉蚜为了有效地获得生存所
需的积温，一般体表面积较大、体色较深。同时在对环境的长期适应过程中，棉蚜成虫
的寿命和产卵量也呈现一定程度的分化。低纬度地区地处热带常年高温，这使得生活在
其中的棉蚜种群经过长期的进化作用，对高温己产生了一定的适应性，因而在高温的条
件下，成虫的寿命较长，产卵量都较大；而高纬度地区的棉蚜，由于对低温的适应，反
而是在低温的条件下，呈现相应的变化。在低纬度地区，由于环境条件适宜，棉蚜终生
行孤雌生殖，不出现有性世代；而在高纬度地区，为了度过寒冷的冬季，棉蚜主要以有
性世代产生的受精卵越冬。可以看出，棉蚜对于不同的生态环境都有其独特的适应对策，
而所有这些生态适应对策，都为其广泛分布，发生危害提供了便利条件。
Cotton aphid Aphis gossypii (Homoptera: Aphididae), an important agricultural pest, its distribution extends from 60 N to 40 S in the world. This insect is capable to survive under different climatic conditions due to its wider adaptability. In summer, this insect reproduces parthenogenetically from generation to generation, after the late autumn it produces sexual aphids to lay eggs to over-winter. This paper presents studies on ecological adaptability of Aphis gossypii under different temperature and photoperiod ranges in the laboratory.
Adults of Aphis gossypii were collected from cotton plants in Ho Chi Minh city of Vietnam (106.42 E, 10.45 N), Beijing (116.5 E, 39.9 N), Shihezi city of Xinjiang Municipality (85 E, 45 N) and Sanya city of Hainan Province (109.5 E, 18.2 N) in 2001 and these adult populations were named as YN, BJ, XJ and HN respectively. Each aphid culture was maintained on cotton seedlings at 22 C, 70%RH, and a photoperiod of 14:10 (L:D). The morphological variation of growth and development and relationship between emergence of sexual morphs and photoperiod (L:D=8:16 and L:D=10:14) at 18 C as well as the life parameters of these populations at different temperatures were evaluated.
In the 1st part of the study with asexual aphids, fifty aphids from each of the 4 populations (i.e. YN, HN, BJ and XJ) were placed at various temperatures (15, 20, 25, 30 and 35 C respectively) maintaining 70%RH, and photoperiod L:D=16:8. Each aphid from respective population was fed on cotton cotyledon in 25ml glass beaker with 10ml-dissolved medium at the bottom. The newly produced aphids were transferred on cotyledon, and the cotton cotyledon was put on the surface of culture medium tightly. The newly produced nymphs were picked out everyday, but the adults were allowed to remain in the same container with replacement of fresh cotyledon and medium as and when required (Leaf Wafer Method). In the experiment, forty newly produced nymphs and the dead adults were all kept. Morphological parameters, such as foreleg femur length, foreleg tibia length, midleg femur length, midleg tibia length, hindleg femur length, hindleg tibia length, body length and body width were studied.
In the 2nd part of the study with sexual aphids (collected from cotton plants), forty 4th instar aphids (G0) were transferred into environment chamber and allowed to breed using LWM. After 24 hours, the G0 adult aphids were removed, and forty new nymphs (G1) from G0 generation were introduced to breed until generation three (G3). The G2 nymphs were kept on feeding, which were produced at the 2nd, 7th, 12th and 17th day separately. During the growth period, the number of apteral and alate aphids was checked and the alate ones were transferred to cotton plants having 3 to 4 leaves to get G3 generation. The cotton plants were enclosed with transparent plastic sleeves, which were covered at the top with gauze. The rates
of development of three types of aphids (1) virginoparae cotton aphid in G3 with reproductive potential, (2) male alate aphid with genetalia without reproductive potential and, (3) female apteral aphid having eggs ] were observed.
In the 3rd part of the study, fifty aphids from each of the 3 populations (i.e. YN, BJ and XJ) were placed at various temperatures (15, 20, 25, 30 and 35 C respectively) maintaining 70%RH, and photoperiod L:D=16:8. Each aphid from respective population was breed with LWM. Ecdysis, reproduction and death of aphid were observed everyday until all aphid adults died.
The results showed that there were significant differences of morphology and color while they developed under various temperatures. Length of foreleg femur, foreleg tibia, midleg femur, midleg tibia, hindleg femur and hindleg tibia, head width and cornicles length of aphid from HN showed significant differences with those from BJ and XJ population. The surface area of both adult and larval aphids gradually increased with decreased temperature. It was found that there were also observed significant differences in surface area among Y
引文
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