温热带地区不同生物型棉蚜对夏寄主的适应性研究
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摘要
棉蚜是农业、畜牧业最主要的世界性害虫之一。在适应不同地区，寄主及其他各种复杂的环境条件的过程中，棉蚜逐渐形成了极为复杂的生活史，寄主专化型和不同的生殖模型。它兼有孤雌生生殖和两性生殖，有研究说前者比后者有更强的进化优势。但是在热带地区生活终年营孤雌生殖的棉蚜是否有足够的进化动力？有自然状态下，寄主转换现象会不会发生？有无取食经验对于成蚜的寄主选择有着什么样的联系？到目前为止，这些问题还没有精确而全面的回答。
目前的研究表明棉蚜至少有两种寄主专化的生物型，一种在棉花上，一种在葫芦科作物上。为弄清两种寄主专化型棉蚜在寄主变化后的适应度的变化，作者进行了以下两个方面的研究：一是测定两种生物型棉蚜在不同作物上的适应能力、偏好性及种群增长率；二是测定有无取食经验的棉蚜在以上方面有无差异。根据成蚜有无取食经验及所在的寄主植物，分为C-WE（有取食经验的棉花种群）、C-NE（无取食经验的棉花种群）、M-WE（有取食经验的甜瓜种群）和M-NE（无取食经验的甜瓜种群）。结果如下：
在转换寄主后适应性试验中，棉花种群被转到甜瓜上时，成蚜寿命为39.83±2.88h（C-WE），62.58±3.50h（C-NE）比转到原寄主棉花的成蚜寿命明显降低，而与转接后无食物的培养皿中的成蚜寿命无明显差异。甜瓜种群的成蚜被转到棉花植株时也有同样的表现。甜瓜种群寿命32±5.28h（M-WE），46.93±2.47h（M-NE）比转到原寄主甜瓜上的寿命明显减少，而与无食物的蚜虫寿命明显差异。甜瓜种群比棉花种群有更长的存活时间，M-NE比M-WE有更长的存活时间，且他们之间均存在着明显差异。有无取食经验对棉花种群的雌成蚜存活时间则无影响。从生殖力来分析，转接到原寄主时，M-NE产蚜量最高（31.44±3.27），与M-WE产蚜量（26.33±2.99）有显著差异。而有无取食经验对棉花种群的产蚜量无影响。从棉花转接至甜瓜、从甜瓜转接至棉花与无食物时成蚜的产蚜量之间无明显差异。
在对两种寄主自然选择试验中，不管是棉花种群还是甜瓜种群，其对原寄主的选择都呈强选择性。有无取食经验对其影响不大。甜瓜种群对原寄主的选择性比棉花种群强。24h与48h后棉花种群对选择原寄主棉花的蚜虫比选择甜瓜的数量之间存在着明显差异，说明其有重新选择的现象。甜瓜种群24h即呈现对原寄主甜瓜的强选择性，48h仍有部分蚜虫从棉花转移到甜瓜，其选择棉花和甜瓜的数量之间皆有明显差异。M-WE比C-WE具备更高的产若蚜能力。M-NE选择甜瓜后若蚜增长速率比M-WE增长更快。
在嗅觉测定试验中，棉花种群和甜瓜种群都更倾向于选择原寄主。C-WE选择新寄主甜瓜的蚜虫数量更多。M-WE在对棉花和水对照进行选择时，对棉花的选择明显增多，且两者差异明显。M-NE在对棉花-水对照的选择中对水对照的选择更多。从蚜虫的选择速度上来分析， C-WE在对棉花-甜瓜进行选择时爬向棉花的时间（258.11±11.55s）比爬向甜瓜的时间（281.52±19.64s）更短。C-NE在对棉花-甜瓜，棉花-水对照，甜瓜-水对照进行选择时，其爬行时间上无显著差异。M-WE在对棉花-甜瓜进行选择时，选择棉花的时间（242.21±11.14s）比选择原寄主甜瓜用的时间（273.6±44.21s）少。M-NE在对棉花-水对照进行选择时，选择棉花的时间（366.43±53.04s）则明显要长一些，而更快地爬向清水（250.35±33.47s）；而在对甜瓜-水对照作选择时，则更快地爬向原寄主甜瓜（281.60±18.24s）。C-WE在对甜瓜-棉花进行选择时，爬向棉花及甜瓜的时间都要比M-WE需要的时间长；M-NE对于棉花-水对照进行选择时，选择棉花所用时间（366.43±53.04s）明显比C-NE所用时间(209.61±2.24s)长，两者之存在明显差异。
在对棉花及10种葫芦科作物的选择试验中，棉花种群的棉蚜对杂交葫芦、小西葫芦表现出强选择性及适应性，而对香瓜、黄瓜、西瓜及甜瓜均不选择；甜瓜种群对甜瓜、香瓜、南瓜、黄西葫芦、大西葫芦、杂交葫芦等均表现较强选择性，而对棉花及小西芦表现不选择。但是，甜瓜种群在小西葫芦上也能产若蚜并存活。从而证明小西葫芦是两种寄主型棉蚜的共同寄主，有可能成为两者相互转换的桥梁寄主。棉花种群的有翅雌成蚜24h后对杂交葫芦的选择性最强，其次为南瓜、棉花、黄西葫芦、小西葫芦、大西葫芦；最后为香瓜、黄瓜、西瓜、冬南瓜及甜瓜。甜瓜型棉蚜而在香瓜、西瓜上的种群增长率最高，与其他各寄主存在统计学意义上的明显差异；其次为甜瓜；再次为南瓜、小西葫芦、冬南瓜；再次为黄瓜、黄西葫芦、大西葫芦及杂交葫芦，在棉花上不能存活。
Aphis gossypii (Glover)(Hemiptera: Aphididae) is one of the most important insect pestsin agriculture and horticulture crops worldwide. In the course of adapting to different areas,hosts and other complicated environmental conditions; it has an extremely complex lifecycle,host specialization and different reproduction models. It has both parthenogenetic and sexualreproduction, and it was said that the former had a strong selective advantage over the latter.But did the cotton aphid have enough evolutionary motivation who only had parthenogeneticreproduction and led an autoecious life cycle in the tropical area? Does host-alternatingphenomenon exist in the nature? And what kind of connection with the feeding experience ofadults of aphid and the host choice to hosts? So far, there have been no surfeint and accurateanswers in the literature.
It was reported that the cotton aphid had at least two specialized biotypes, one on cotton,and the other on cucurbit host plants. In order to determine the differences of fitness of thetwo biotypes of cotton aphid which belong to two specializations when the hosts changed, Iinitiated the studies with two objectives:(1) to investigate the adaptation ability on differenthost plants, and the preference and finite increase rates on10species of Cucurbtaceae withthe winged adults which were the natural form in the field; and (2)to determine the differencesbetween the adults which had feeding experience and without experience. According to theadults with or without feeding experience on cotton and melon, we divided the experimentaladults into4groups: C-WE, C-NE, M-WE and M-NE.The results were summarized below:
In the experiment of host adaptation-no choice test, the longevity of the cotton biotypewith feeding experience(C-WE) is much shorter(39.83±2.88h) than the ones withoutexperience(C-NE62.58±3.50h) when they were transferred from cotton to melon plants. Andthat result had no statistic significant differences between the ones which had no food. Same,the longevity of the adults from melon (32±5.28h M-WE),(46.93±2.47h M-NE) were alsoshorter than the ones who were transferred to original host plants. The longevity of adultsfrom melon were longer than the ones from cotton. The group M-NE had longer survive time than the group with experience(M-WE). But the ones from cotton had no difference betweenthe ones with experience and without experience. In the same time, the offspring of the groupM-NE was highest（31.44±3.27）, and this result had statistic significant difference betweenM-WE (26.33±2.99). Otherwise, there were no significant difference between the ones withand without feeding experience from cotton.
In the experiment of host adaptation-choice test, the adults from two hosts both hadstrong preference to the original host plants. And the feeding experience had no effect on thechoice to hosts. The number of adults on cotton from cotton group after48h was much morethan24h after adult introduction. That revealed that the adults had re-selection phenomenon.Also, the offspring of the group M-WE was higher than the group C-WE. The increasing rateof the group from melon was also higher than the ones from cotton. The group M-NE hadhigher increasing rate when they chose the same host than the group M-WE.
In the olfactory experiment, the original hosts obviously affected the choice to the adults inthe re-selection process. Both of the two groups preferred to choose the original host plants,indicating that the plant volatiles did work in the selection process of the host plants. Moreadults of C-WE chose the new host-melon. And more adults of C-WE chose melon whenthey had to choose from melon and water, and melon and cotton. Interestingly, more adults ofM-NE preferred water rather than cotton plants. The time of adults of C-WE which crawlingto cotton (258.11±11.55s)was shorter than to melon (281.52±19.64s). But the time of theadults of C-NE was not different when they crawling to hosts or water. The adults of M-WEhad shorter time to choose cotton(242.21±11.14s) than to the original host melon(273.6±44.21s) when they chose cotton and melon plants. And the group of M-NE spentlonger time(366.43±53.04s) to choose cotton plants than to water(250.35±33.47s) when theychose cotton and the control water. Also, this group used shorter time to choose original hostmelon(281.60±18.24s) than the time to water.
Comparison between the two groups, the adults of C-WE spent longer crawling time thanthe ones of M-WE. The adults of M-NE used significant longer time (366.43±53.04s)crawling to cotton plants than the ones of C-NE (209.61±2.24s).
In the test of the preference of the aphids to different summer hosts including cotton and10kinds of Cucurbtaceae plants, we determined the preference and finite increase rates of thecotton and melon aphids on10hosts of Cucurbtaceae with the winged adults. The adults fromcotton significantly preferred hybrid squash and zucchini to other host plants in the first day.And the ones from melon chose most of the hosts including melon, cantaloupe, pumpkin,yellow squash, tornado and hybrid squash, but not cotton and zucchini. But, even the aphids from melon did not prefer zucchini; however, they could survive on this host well. We thinkthat zucchini should be the mutual host of the two aphid biotypes, serving as a bridge host ofthe two aphid biotypes.
目前的研究表明棉蚜至少有两种寄主专化的生物型，一种在棉花上，一种在葫芦科作物上。为弄清两种寄主专化型棉蚜在寄主变化后的适应度的变化，作者进行了以下两个方面的研究：一是测定两种生物型棉蚜在不同作物上的适应能力、偏好性及种群增长率；二是测定有无取食经验的棉蚜在以上方面有无差异。根据成蚜有无取食经验及所在的寄主植物，分为C-WE（有取食经验的棉花种群）、C-NE（无取食经验的棉花种群）、M-WE（有取食经验的甜瓜种群）和M-NE（无取食经验的甜瓜种群）。结果如下：
在转换寄主后适应性试验中，棉花种群被转到甜瓜上时，成蚜寿命为39.83±2.88h（C-WE），62.58±3.50h（C-NE）比转到原寄主棉花的成蚜寿命明显降低，而与转接后无食物的培养皿中的成蚜寿命无明显差异。甜瓜种群的成蚜被转到棉花植株时也有同样的表现。甜瓜种群寿命32±5.28h（M-WE），46.93±2.47h（M-NE）比转到原寄主甜瓜上的寿命明显减少，而与无食物的蚜虫寿命明显差异。甜瓜种群比棉花种群有更长的存活时间，M-NE比M-WE有更长的存活时间，且他们之间均存在着明显差异。有无取食经验对棉花种群的雌成蚜存活时间则无影响。从生殖力来分析，转接到原寄主时，M-NE产蚜量最高（31.44±3.27），与M-WE产蚜量（26.33±2.99）有显著差异。而有无取食经验对棉花种群的产蚜量无影响。从棉花转接至甜瓜、从甜瓜转接至棉花与无食物时成蚜的产蚜量之间无明显差异。
在对两种寄主自然选择试验中，不管是棉花种群还是甜瓜种群，其对原寄主的选择都呈强选择性。有无取食经验对其影响不大。甜瓜种群对原寄主的选择性比棉花种群强。24h与48h后棉花种群对选择原寄主棉花的蚜虫比选择甜瓜的数量之间存在着明显差异，说明其有重新选择的现象。甜瓜种群24h即呈现对原寄主甜瓜的强选择性，48h仍有部分蚜虫从棉花转移到甜瓜，其选择棉花和甜瓜的数量之间皆有明显差异。M-WE比C-WE具备更高的产若蚜能力。M-NE选择甜瓜后若蚜增长速率比M-WE增长更快。
在嗅觉测定试验中，棉花种群和甜瓜种群都更倾向于选择原寄主。C-WE选择新寄主甜瓜的蚜虫数量更多。M-WE在对棉花和水对照进行选择时，对棉花的选择明显增多，且两者差异明显。M-NE在对棉花-水对照的选择中对水对照的选择更多。从蚜虫的选择速度上来分析， C-WE在对棉花-甜瓜进行选择时爬向棉花的时间（258.11±11.55s）比爬向甜瓜的时间（281.52±19.64s）更短。C-NE在对棉花-甜瓜，棉花-水对照，甜瓜-水对照进行选择时，其爬行时间上无显著差异。M-WE在对棉花-甜瓜进行选择时，选择棉花的时间（242.21±11.14s）比选择原寄主甜瓜用的时间（273.6±44.21s）少。M-NE在对棉花-水对照进行选择时，选择棉花的时间（366.43±53.04s）则明显要长一些，而更快地爬向清水（250.35±33.47s）；而在对甜瓜-水对照作选择时，则更快地爬向原寄主甜瓜（281.60±18.24s）。C-WE在对甜瓜-棉花进行选择时，爬向棉花及甜瓜的时间都要比M-WE需要的时间长；M-NE对于棉花-水对照进行选择时，选择棉花所用时间（366.43±53.04s）明显比C-NE所用时间(209.61±2.24s)长，两者之存在明显差异。
在对棉花及10种葫芦科作物的选择试验中，棉花种群的棉蚜对杂交葫芦、小西葫芦表现出强选择性及适应性，而对香瓜、黄瓜、西瓜及甜瓜均不选择；甜瓜种群对甜瓜、香瓜、南瓜、黄西葫芦、大西葫芦、杂交葫芦等均表现较强选择性，而对棉花及小西芦表现不选择。但是，甜瓜种群在小西葫芦上也能产若蚜并存活。从而证明小西葫芦是两种寄主型棉蚜的共同寄主，有可能成为两者相互转换的桥梁寄主。棉花种群的有翅雌成蚜24h后对杂交葫芦的选择性最强，其次为南瓜、棉花、黄西葫芦、小西葫芦、大西葫芦；最后为香瓜、黄瓜、西瓜、冬南瓜及甜瓜。甜瓜型棉蚜而在香瓜、西瓜上的种群增长率最高，与其他各寄主存在统计学意义上的明显差异；其次为甜瓜；再次为南瓜、小西葫芦、冬南瓜；再次为黄瓜、黄西葫芦、大西葫芦及杂交葫芦，在棉花上不能存活。
Aphis gossypii (Glover)(Hemiptera: Aphididae) is one of the most important insect pestsin agriculture and horticulture crops worldwide. In the course of adapting to different areas,hosts and other complicated environmental conditions; it has an extremely complex lifecycle,host specialization and different reproduction models. It has both parthenogenetic and sexualreproduction, and it was said that the former had a strong selective advantage over the latter.But did the cotton aphid have enough evolutionary motivation who only had parthenogeneticreproduction and led an autoecious life cycle in the tropical area? Does host-alternatingphenomenon exist in the nature? And what kind of connection with the feeding experience ofadults of aphid and the host choice to hosts? So far, there have been no surfeint and accurateanswers in the literature.
It was reported that the cotton aphid had at least two specialized biotypes, one on cotton,and the other on cucurbit host plants. In order to determine the differences of fitness of thetwo biotypes of cotton aphid which belong to two specializations when the hosts changed, Iinitiated the studies with two objectives:(1) to investigate the adaptation ability on differenthost plants, and the preference and finite increase rates on10species of Cucurbtaceae withthe winged adults which were the natural form in the field; and (2)to determine the differencesbetween the adults which had feeding experience and without experience. According to theadults with or without feeding experience on cotton and melon, we divided the experimentaladults into4groups: C-WE, C-NE, M-WE and M-NE.The results were summarized below:
In the experiment of host adaptation-no choice test, the longevity of the cotton biotypewith feeding experience(C-WE) is much shorter(39.83±2.88h) than the ones withoutexperience(C-NE62.58±3.50h) when they were transferred from cotton to melon plants. Andthat result had no statistic significant differences between the ones which had no food. Same,the longevity of the adults from melon (32±5.28h M-WE),(46.93±2.47h M-NE) were alsoshorter than the ones who were transferred to original host plants. The longevity of adultsfrom melon were longer than the ones from cotton. The group M-NE had longer survive time than the group with experience(M-WE). But the ones from cotton had no difference betweenthe ones with experience and without experience. In the same time, the offspring of the groupM-NE was highest（31.44±3.27）, and this result had statistic significant difference betweenM-WE (26.33±2.99). Otherwise, there were no significant difference between the ones withand without feeding experience from cotton.
In the experiment of host adaptation-choice test, the adults from two hosts both hadstrong preference to the original host plants. And the feeding experience had no effect on thechoice to hosts. The number of adults on cotton from cotton group after48h was much morethan24h after adult introduction. That revealed that the adults had re-selection phenomenon.Also, the offspring of the group M-WE was higher than the group C-WE. The increasing rateof the group from melon was also higher than the ones from cotton. The group M-NE hadhigher increasing rate when they chose the same host than the group M-WE.
In the olfactory experiment, the original hosts obviously affected the choice to the adults inthe re-selection process. Both of the two groups preferred to choose the original host plants,indicating that the plant volatiles did work in the selection process of the host plants. Moreadults of C-WE chose the new host-melon. And more adults of C-WE chose melon whenthey had to choose from melon and water, and melon and cotton. Interestingly, more adults ofM-NE preferred water rather than cotton plants. The time of adults of C-WE which crawlingto cotton (258.11±11.55s)was shorter than to melon (281.52±19.64s). But the time of theadults of C-NE was not different when they crawling to hosts or water. The adults of M-WEhad shorter time to choose cotton(242.21±11.14s) than to the original host melon(273.6±44.21s) when they chose cotton and melon plants. And the group of M-NE spentlonger time(366.43±53.04s) to choose cotton plants than to water(250.35±33.47s) when theychose cotton and the control water. Also, this group used shorter time to choose original hostmelon(281.60±18.24s) than the time to water.
Comparison between the two groups, the adults of C-WE spent longer crawling time thanthe ones of M-WE. The adults of M-NE used significant longer time (366.43±53.04s)crawling to cotton plants than the ones of C-NE (209.61±2.24s).
In the test of the preference of the aphids to different summer hosts including cotton and10kinds of Cucurbtaceae plants, we determined the preference and finite increase rates of thecotton and melon aphids on10hosts of Cucurbtaceae with the winged adults. The adults fromcotton significantly preferred hybrid squash and zucchini to other host plants in the first day.And the ones from melon chose most of the hosts including melon, cantaloupe, pumpkin,yellow squash, tornado and hybrid squash, but not cotton and zucchini. But, even the aphids from melon did not prefer zucchini; however, they could survive on this host well. We thinkthat zucchini should be the mutual host of the two aphid biotypes, serving as a bridge host ofthe two aphid biotypes.
引文
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