寄主体型和日龄对广大腿小蜂产卵选择和后代发育适合度的影响
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摘要
产卵选择与后代发育适合度之间的相关性是进化生态学的主要科学问题之一。在寄生蜂中,一般认为寄主体型大小是寄生蜂产卵决策的主要依据,即“寄主大小-品质’假说。但寄主日龄也可能影响寄生蜂的决策,因为寄主日龄与其生理变化密切相关,从而影响子代蜂的发育。检验“寄主大小-品质”假说常用体型大小不同的异种寄主或体型大小变异较大的同种寄主,前者存在遗传差异难以克服的问题,后者则依赖于那些种内个体间体型大小存在很大差异的寄主。为了克服上述问题,本论文研究以广大腿小蜂(Brachymeria lasus)及其寄主菜粉蝶(Pieris rapae)蛹为材料,采用饥饿方法获得体型差异较大的同种寄主蛹供寄生,观察寄生蜂的产卵行为及其后代发育适合度表现,以检验该假说。同时,研究了寄主蛹龄等对产卵选择及后代适合度的影响。研究获得以下主要结果。
1.菜粉蝶蛹体型大小对广大腿小蜂寄生选择及后代发育的影响
为检验“寄主大小-品质”假说,采取饥饿方法处理菜粉蝶4龄幼虫,以获得体型大小(用鲜重表示)差异大于同一自然地理种群内个体间差异的寄主蛹,然后供广大腿小蜂寄生,观察寄生选择行为,以及子代蜂发育历期、性比和体型大小等适应度相关特征。研究结果表明：(i)广大腿小蜂搜寻寄主的时间以及寄生完毕后的整饰时间与寄主体型大小无关,产卵时间与寄主体型大小呈正相关关系,休息和爬行时间与寄主体型大小呈负相关关系；(ii)非选择性和选择性试验表明,广大腿小蜂显著偏好体型较大的寄主蛹,而且在体型较大的蛹内产雌性后代的概率更大,寄主蛹体型大小与子代蜂体型大小存在显著的正相关性,但与子代蜂发育历期无关。上述研究结果说明,广大腿小蜂对菜粉蝶蛹的寄生选择与后代发育表现,符合“寄主大小-品质”假说的预测。
2.菜粉蝶蛹龄对广大腿小蜂寄生选择及后代发育的影响
为了探明寄主菜粉蝶蛹龄对广大腿小蜂产卵选择和后代发育的影响,采用饥饿法处理菜粉蝶4龄幼虫,从而获得不同体型大小的菜粉蝶蛹。将饥饿处理获得的蛹(体型较小)和对照蛹(体型较大)分为3个日龄组：1d、3d和5d,供寄生蜂寄生。研究结果表明：(i)菜粉蝶蛹龄和体型大小之间不存在互作关系；(ii)菜粉蝶蛹龄对广大腿小蜂分配雌性的概率没有影响；仅与菜粉蝶蛹体型有关；(iii)子代雌蜂发育历期与寄主蛹龄存在微弱但显著的相关性,但子代雄蜂的发育历期、雌雄蜂的寿命及体型大小等发育适应度表现均不受寄主蛹龄的显著影响。上述研究结果说明,寄主蛹龄不影响广大腿小蜂的产卵选择,但对子代蜂发育有一定的影响。
3.学习在广大腿小蜂寄生选择行为中的作用
为探究寄生经历是否影响广大腿小蜂后续产卵选择行为,对2个不同的观测变量—选择频次和寄主识别期进行了实验观察和分析。利用选择性实验方法,在盆栽植物上接种寄主菜粉蝶蛹(饥饿处理蛹和对照蛹),采取行为连续观察法记录寄生蜂发生第1次和第2次产卵行为的时间和对象,研究结果表明：(i)具有寄生经历的广大腿小蜂显著偏好寄生体型较大的菜粉蝶蛹；(ii)但寄生经历对广大腿小蜂的寄主识别期没有显著影响,由此说明,寄主识别期不受寄生经历的影响。
Relationship between parasitization preference and progeny performance is one of central themes in evolutionary ecology. For most parasitoids, host body size is usually regarded as the main accordance in oviposition strategy, which is known in "host size-quality"hypothesis. However, host age also has impact on oviposition strategy of parasitoids, for its changes will be along with physiological transformation which will affect the developmental performance of offspring parasitoids. So far, there have been two methods to test "host size-quality" hypothesis, one is using different host species with wide body size variation; the other is using the same host specie with broad range in body size. However, the former had unconquerable problems in inheritance differences; the latter relyed on the variation in body size in the same host specie. So in this thesis study, to test the "host size-quality" hypothesis,4th instar larvae of P. rapae were manipulated by starvation whose pupae were exposed to parasitism by B. lasus. Then the oviposition behaviors of mother parasitoids; the development performances of offspring parasitoids were recorded. Meanwhile, host age was tested as another impact factor. The main results are as followed below.
1. Influence of host pupa weight on parasitization selection and offspring developmental performance of Brachymeria lasus
To test the "host size-quality" hypothesis,4th instar larvae of P. rapae were manipulated by starvation regime to obtain a wide range of host body size, and these pupae were individually exposed to parasitism by the solitary parasitoid B. lasus. Then the behaviors of parasitization selection of mother parasitoids; development time, sex ratio and body size of offspring parasitoids were recorded. The results showed that (ⅰ) searching time and preening time were noncorrelated with host body size; ovipositioning time was correlated with host size positively while the resting and creeping time was correlated with that negatively; (ii) the results of non-choice and choice experiment suggested that parasitoids prefer to lay eggs in larger host pupae, and more females are produced from larger host pupae. The body size of male and female offsprings were increased with host body size in a linear relationship, yet the development time remained constant regardless of host body size. The results indicate that the influence of host pupa weight on parasitization decision and offspring developmental performance of B. lasus are in accordance with the predictions of "host size-quality" hypothesis.
2. Effects of host pupa age on parasitization decision and offspring developmental performance of Brachymeria lasus
To investigate the effects of host pupa age on parasitization decision and offspring developmental performances of the parasitoids, larvae of P. rapae were manipulated by starvation regime to obtain a wide range of host body size. Those pupae at age of either 1-day,3-day, and 5-day old were individually exposed to parasitism. The results showed that (i)there was no interaction between host body size and host age; (ii) host age did not have impact on probability of female offsprings, which was only affected by host body size,; (iii) there was a weak yet significant relationship between female development time and host age, but other developmental performances (including male development time、offspring lifespan and body size)were not influenced by host age. The results suggest that host pupa age does not affect the parasitization decision of B. lasus, but it has impact on offspring development to some extent.
3. The effect of learning in parasitization selection of B. lasus
To assess whether parasitization experience would affect the latter oviposition behaviors of B. lasus or not, two observation values—selection frequency and host recognition time were selected to observe and analyze. Under choice condition, two host pupae with starvation treated and another two without were put on a potted cabbage to observe the first and succecive ovipostion of a single parasitoid. The results were followed (i)the Chiqs-test of selection frequency showed that those parasitoids with parasitization experience would prefer to lay eggs on larger host pupae significantly; (ii)however, parasitization experience did not influence the parasitzation preference of B. lasus when it was evaluated by host recognition time.
1.菜粉蝶蛹体型大小对广大腿小蜂寄生选择及后代发育的影响
为检验“寄主大小-品质”假说,采取饥饿方法处理菜粉蝶4龄幼虫,以获得体型大小(用鲜重表示)差异大于同一自然地理种群内个体间差异的寄主蛹,然后供广大腿小蜂寄生,观察寄生选择行为,以及子代蜂发育历期、性比和体型大小等适应度相关特征。研究结果表明：(i)广大腿小蜂搜寻寄主的时间以及寄生完毕后的整饰时间与寄主体型大小无关,产卵时间与寄主体型大小呈正相关关系,休息和爬行时间与寄主体型大小呈负相关关系；(ii)非选择性和选择性试验表明,广大腿小蜂显著偏好体型较大的寄主蛹,而且在体型较大的蛹内产雌性后代的概率更大,寄主蛹体型大小与子代蜂体型大小存在显著的正相关性,但与子代蜂发育历期无关。上述研究结果说明,广大腿小蜂对菜粉蝶蛹的寄生选择与后代发育表现,符合“寄主大小-品质”假说的预测。
2.菜粉蝶蛹龄对广大腿小蜂寄生选择及后代发育的影响
为了探明寄主菜粉蝶蛹龄对广大腿小蜂产卵选择和后代发育的影响,采用饥饿法处理菜粉蝶4龄幼虫,从而获得不同体型大小的菜粉蝶蛹。将饥饿处理获得的蛹(体型较小)和对照蛹(体型较大)分为3个日龄组：1d、3d和5d,供寄生蜂寄生。研究结果表明：(i)菜粉蝶蛹龄和体型大小之间不存在互作关系；(ii)菜粉蝶蛹龄对广大腿小蜂分配雌性的概率没有影响；仅与菜粉蝶蛹体型有关；(iii)子代雌蜂发育历期与寄主蛹龄存在微弱但显著的相关性,但子代雄蜂的发育历期、雌雄蜂的寿命及体型大小等发育适应度表现均不受寄主蛹龄的显著影响。上述研究结果说明,寄主蛹龄不影响广大腿小蜂的产卵选择,但对子代蜂发育有一定的影响。
3.学习在广大腿小蜂寄生选择行为中的作用
为探究寄生经历是否影响广大腿小蜂后续产卵选择行为,对2个不同的观测变量—选择频次和寄主识别期进行了实验观察和分析。利用选择性实验方法,在盆栽植物上接种寄主菜粉蝶蛹(饥饿处理蛹和对照蛹),采取行为连续观察法记录寄生蜂发生第1次和第2次产卵行为的时间和对象,研究结果表明：(i)具有寄生经历的广大腿小蜂显著偏好寄生体型较大的菜粉蝶蛹；(ii)但寄生经历对广大腿小蜂的寄主识别期没有显著影响,由此说明,寄主识别期不受寄生经历的影响。
Relationship between parasitization preference and progeny performance is one of central themes in evolutionary ecology. For most parasitoids, host body size is usually regarded as the main accordance in oviposition strategy, which is known in "host size-quality"hypothesis. However, host age also has impact on oviposition strategy of parasitoids, for its changes will be along with physiological transformation which will affect the developmental performance of offspring parasitoids. So far, there have been two methods to test "host size-quality" hypothesis, one is using different host species with wide body size variation; the other is using the same host specie with broad range in body size. However, the former had unconquerable problems in inheritance differences; the latter relyed on the variation in body size in the same host specie. So in this thesis study, to test the "host size-quality" hypothesis,4th instar larvae of P. rapae were manipulated by starvation whose pupae were exposed to parasitism by B. lasus. Then the oviposition behaviors of mother parasitoids; the development performances of offspring parasitoids were recorded. Meanwhile, host age was tested as another impact factor. The main results are as followed below.
1. Influence of host pupa weight on parasitization selection and offspring developmental performance of Brachymeria lasus
To test the "host size-quality" hypothesis,4th instar larvae of P. rapae were manipulated by starvation regime to obtain a wide range of host body size, and these pupae were individually exposed to parasitism by the solitary parasitoid B. lasus. Then the behaviors of parasitization selection of mother parasitoids; development time, sex ratio and body size of offspring parasitoids were recorded. The results showed that (ⅰ) searching time and preening time were noncorrelated with host body size; ovipositioning time was correlated with host size positively while the resting and creeping time was correlated with that negatively; (ii) the results of non-choice and choice experiment suggested that parasitoids prefer to lay eggs in larger host pupae, and more females are produced from larger host pupae. The body size of male and female offsprings were increased with host body size in a linear relationship, yet the development time remained constant regardless of host body size. The results indicate that the influence of host pupa weight on parasitization decision and offspring developmental performance of B. lasus are in accordance with the predictions of "host size-quality" hypothesis.
2. Effects of host pupa age on parasitization decision and offspring developmental performance of Brachymeria lasus
To investigate the effects of host pupa age on parasitization decision and offspring developmental performances of the parasitoids, larvae of P. rapae were manipulated by starvation regime to obtain a wide range of host body size. Those pupae at age of either 1-day,3-day, and 5-day old were individually exposed to parasitism. The results showed that (i)there was no interaction between host body size and host age; (ii) host age did not have impact on probability of female offsprings, which was only affected by host body size,; (iii) there was a weak yet significant relationship between female development time and host age, but other developmental performances (including male development time、offspring lifespan and body size)were not influenced by host age. The results suggest that host pupa age does not affect the parasitization decision of B. lasus, but it has impact on offspring development to some extent.
3. The effect of learning in parasitization selection of B. lasus
To assess whether parasitization experience would affect the latter oviposition behaviors of B. lasus or not, two observation values—selection frequency and host recognition time were selected to observe and analyze. Under choice condition, two host pupae with starvation treated and another two without were put on a potted cabbage to observe the first and succecive ovipostion of a single parasitoid. The results were followed (i)the Chiqs-test of selection frequency showed that those parasitoids with parasitization experience would prefer to lay eggs on larger host pupae significantly; (ii)however, parasitization experience did not influence the parasitzation preference of B. lasus when it was evaluated by host recognition time.
引文
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