菜粉蝶蛹质量对蝶蛹金小蜂产卵决策的影响
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摘要
寄生蜂是检验行为生态学和进化生态学假说的理想模式生物。寄生蜂对寄主的选择是寄主质量和寄主价值共同作用的过程,寄主日龄和寄主体型大小是决定寄主质量的两个主要变量,在一定程度上决定寄生蜂后代数量、性比及后代适合度。迄今,“寄主大小-质量”假说在抑性寄生蜂(产卵后寄主停止发育)中获得了大量研究的支持,但寄主日龄也可能影响这一模型的预测,因为寄主日龄越高,营养越差。本论文研究以群集寄生性蝶蛹金小蜂Pteromalus puparum及其寄主菜粉蝶Pieris rapae蛹为对象,为了克服检验“寄主大小-质量”模型的传统方法(用体型大小不同的异种寄主)的缺陷—难以避免寄主种间遗传差异的干扰,用饥饿方法获得体型大小差异较大的同种寄主蛹,观察寄生蜂的产卵行为及其后代发育适合度表现,以求更精确地检验该模型；另外,还观察了寄生蜂不同日龄、不同蛹龄等变量对产卵策略及其后代适合度的影响。研究获得以下主要结果。
1.菜粉蝶蛹体型大小对蝶蛹金小蜂产卵策略的影响
为检验“寄主大小-质量”假说,采取饥饿方法处理菜粉蝶Pieris rapae高龄幼虫,以获得体型大小(用重量表示)差异大于同一自然地理种群内个体间差异的寄主蛹,然后供群集蝶蛹金小蜂寄生,观察寄生蜂在不同大小寄主蛹内的后代窝卵数、性比以及体型大小。结果表明,蝶蛹金小蜂1次攻击产出的后代窝卵数随寄主蛹重量增加而显著增多,蛹中平均出蜂314.97头·g-1。蝶蛹金小蜂后代的雌性比例随寄主蛹重量的增大而显著提高,后代雌蜂和雄蜂的体型大小(后足胫节长度)均随寄主蛹重量的增大而显著增大。可见,蝶蛹金小蜂母蜂能够根据寄主蛹重量来调整后代窝卵大小和性比,以使后代适应度最大化。研究结果支持“寄主大小-质量”假说。
2.寄生蜂日龄对蝶蛹金小蜂产卵策略的影响
为摸清寄生蜂日龄对产卵策略的影响,采取饥饿的方法处理菜粉蝶Pieris rapae高龄幼虫,以获得体型大小(用体质量表示)差异显著大于同一自然地理种群内个体间差异程度的寄主蛹,然后供不同日龄(5d、10d、20d和30d)群集蝶蛹金小蜂寄生,观察在不同大小寄主蛹内的后代窝卵数、性比及其体型大小(用后足胫节长度表示)。结果表明,蝶蛹金小蜂日龄显著影响其产卵决策。随着蜂日龄的增加,(ⅰ)后代窝卵数呈明显下降趋势,5d寄生蜂后代平均数量为49.14头,而30d寄生蜂后代平均数量仅为10.84头；(ⅱ)后代雌性比例上升,5d寄生蜂后代平均雌性比例为0.91,而30d寄生蜂为0.95；(ⅲ)后代体型大小总体上呈上升趋势,5d寄生蜂后代雌蜂的平均后足胫节长度为0.68mm,雄蜂后代平均为0.56mm,而30d寄生蜂后代雌蜂的后足胫节长度为0.76mm,雄蜂后代平均为0.61mm。由此推测,蝶蛹金小蜂雌蜂能够通过调整后代窝卵大小和性比,获得最大适应度。
3.菜粉蝶蛹日龄对蝶蛹金小蜂产卵策略的影响
为了探明寄主菜粉蝶蛹日龄对蝶蛹金小蜂产卵决策的影响,采用饥饿法处理菜粉蝶末龄幼虫来获得不同大小的菜粉蝶蛹,然后将不同大小的菜粉蝶蛹随机分组,置于25℃、RH60%～80%、L：D=10：14 h光照培养箱中放置以获得不同日龄的菜粉蝶蛹,供群集蝶蛹金小蜂寄生。研究结果表明,(ⅰ)一次攻击产生的平均后代窝卵数总体上随寄主蛹日龄的增大而显著降低；(ⅱ)后代平均雌性比例随寄主蛹龄的增大而呈下降趋势；(ⅲ)蝶蛹金小蜂后代平均个体大小(用后足胫节长度表示)在前5个日龄(1d、2d、3d、4d和5d)之间变化不明显,而寄生6d蛹的后代蜂后足胫节长度明显变小,而且出蜂率明显降低,仅40%。由此推断,寄主蛹龄是影响蝶蛹金小蜂产卵决策的重要变量之一,所以,用“寄主大小-质量”假说解释抑性寄生蜂产卵策略时,须同时考虑寄主日龄的影响。
4.结论
本论文研究表明,除了寄主蛹体型大小外,寄主蛹日龄以及寄生蜂日龄也显著影响蝶蛹金小蜂的产卵决策。其他潜在因素(如抱卵量、身体营养状态、被捕食和寄生蜂风险等)对蝶蛹金小蜂产卵决策的影响仍有待于进一步研究。
Parasiotids are ideal model systems for testing behavioral and evolutionary hypotheses. Host choice is realized from interaction between host quality and host value in parasitoids. Host age and body size are the two main variables that constitute host quality, as manifested by fitness correlates of offspring parasitoids including clutch size, sex ratio, and body size. As of this time, the "host size-quality" hypothesis has been substantiated by numerous empirical studies of idiobiontic parasitoids. However, host age can influence the prediction of the hypothesis in idiobionts, as host quality may covary with host age. This thesis study, by using gregarious parasitoid Pteromalus puparum and host pupa of Pieris rapae as a model system, was intended to test the host size-quality hypothesis by manipulating host pupa size through starvation of larvae, host age, and parasitoid age as may affect oviposition decision in clutch size and sex ratio. The main results are as followed below.
1. Influence of host pupa weight on oviposition strategy of Pteromalus puparum
To test the "host size-quality"hypothesis, larvae of P. rapae were manipulated by starvation regime to obtain a wide range of body sizeofhost pupa, and then these puae were individually exposed to parasitism by the gregarious parasitoid.After a single attack the pupae were observed for emergence of offspring parasitoids. The results showed that the offspring production was linearly increased with increasing host pupa mass at oviposition, being 314.97 individuals per gram of pupa mass on average. The proportion of offspring females and the body size (hind tibia length) of offspring males and females also increased with increasing host pupa mass. The results suggest that P. puparum can allocate its offspring number and sex ratio, so as to maximize its fitness, basing on assessment of pupaquality. The reproductive strategy of P. puparum and related factors were discussed.
2. Effects of parasitoid age on oviposition strategy of Pteromalus puparum
To investigate how parasitoid age affects oviposition decision as predicted by the host size-quality hypothesis, parasitoids at either of 5-day,10-day,20-day, or 30-day old were provided with host pupae of different body sizes as obtained by the larval starvation. The results shown that parasitoid age significantly affect oviposition decision in P. puparum. With increase of parasitoid age, (ⅰ) clutch size obviously declined, with 49.14 (mean±se) from 5-day old mother wasps and 10.84from the 30-day old; (ⅱ) female increased in offspring, with 0.91 from 5-day old mother wasps and 0.95from the 30-day old; (ⅲ) body size enlarged slightly with 0.68mm in hind tibia length of females and 0.56mm of males from 5-day old mother wasps, and with 0.76mm of females and 0.61mm of males from 30-day old mother wasps. It is suggested that P. puparum might adjust oviposition decision in response to its own age.
3. Effects of host pupa age on oviposition strategy of Pteromalus puparum
To investigate effects of host pupa age on oviposition decision and offspring developmental performances of the parasitoid, host pupae at age of either 1-day,2-day, 3-day,4-day,5-day, and 6-day old were individually exposed to parasitism. The results shown that (ⅰ) clutch size decreased with host age; (2) female proportion declined with host age; (ⅲ) body size was not affected by host age from 1-day through to 5-day, but reduced from attacking 6-day old pupa with much reduced survival by 40%. It is suggested that host pupa age is an important variable in oviposition strategy of Pteromalus puparum, and therefore may confound predictions of the host-size-quality hypothesis.
4. Conclusions
The thesis study indicates that host pupa size is a main variable that measures host quality by gregarious Pteromalus puparum, but host age and mother wasp age are also important variables that affect host quality as perceived by the parasitoid as well. In addition to these variables, other potential traites, such as egg load, nutrition state, risks of predation and parasitism, etc may influence ovipisiton strategy, which are remained for further studies before fully understanding of oviposition strategy.
1.菜粉蝶蛹体型大小对蝶蛹金小蜂产卵策略的影响
为检验“寄主大小-质量”假说,采取饥饿方法处理菜粉蝶Pieris rapae高龄幼虫,以获得体型大小(用重量表示)差异大于同一自然地理种群内个体间差异的寄主蛹,然后供群集蝶蛹金小蜂寄生,观察寄生蜂在不同大小寄主蛹内的后代窝卵数、性比以及体型大小。结果表明,蝶蛹金小蜂1次攻击产出的后代窝卵数随寄主蛹重量增加而显著增多,蛹中平均出蜂314.97头·g-1。蝶蛹金小蜂后代的雌性比例随寄主蛹重量的增大而显著提高,后代雌蜂和雄蜂的体型大小(后足胫节长度)均随寄主蛹重量的增大而显著增大。可见,蝶蛹金小蜂母蜂能够根据寄主蛹重量来调整后代窝卵大小和性比,以使后代适应度最大化。研究结果支持“寄主大小-质量”假说。
2.寄生蜂日龄对蝶蛹金小蜂产卵策略的影响
为摸清寄生蜂日龄对产卵策略的影响,采取饥饿的方法处理菜粉蝶Pieris rapae高龄幼虫,以获得体型大小(用体质量表示)差异显著大于同一自然地理种群内个体间差异程度的寄主蛹,然后供不同日龄(5d、10d、20d和30d)群集蝶蛹金小蜂寄生,观察在不同大小寄主蛹内的后代窝卵数、性比及其体型大小(用后足胫节长度表示)。结果表明,蝶蛹金小蜂日龄显著影响其产卵决策。随着蜂日龄的增加,(ⅰ)后代窝卵数呈明显下降趋势,5d寄生蜂后代平均数量为49.14头,而30d寄生蜂后代平均数量仅为10.84头；(ⅱ)后代雌性比例上升,5d寄生蜂后代平均雌性比例为0.91,而30d寄生蜂为0.95；(ⅲ)后代体型大小总体上呈上升趋势,5d寄生蜂后代雌蜂的平均后足胫节长度为0.68mm,雄蜂后代平均为0.56mm,而30d寄生蜂后代雌蜂的后足胫节长度为0.76mm,雄蜂后代平均为0.61mm。由此推测,蝶蛹金小蜂雌蜂能够通过调整后代窝卵大小和性比,获得最大适应度。
3.菜粉蝶蛹日龄对蝶蛹金小蜂产卵策略的影响
为了探明寄主菜粉蝶蛹日龄对蝶蛹金小蜂产卵决策的影响,采用饥饿法处理菜粉蝶末龄幼虫来获得不同大小的菜粉蝶蛹,然后将不同大小的菜粉蝶蛹随机分组,置于25℃、RH60%～80%、L：D=10：14 h光照培养箱中放置以获得不同日龄的菜粉蝶蛹,供群集蝶蛹金小蜂寄生。研究结果表明,(ⅰ)一次攻击产生的平均后代窝卵数总体上随寄主蛹日龄的增大而显著降低；(ⅱ)后代平均雌性比例随寄主蛹龄的增大而呈下降趋势；(ⅲ)蝶蛹金小蜂后代平均个体大小(用后足胫节长度表示)在前5个日龄(1d、2d、3d、4d和5d)之间变化不明显,而寄生6d蛹的后代蜂后足胫节长度明显变小,而且出蜂率明显降低,仅40%。由此推断,寄主蛹龄是影响蝶蛹金小蜂产卵决策的重要变量之一,所以,用“寄主大小-质量”假说解释抑性寄生蜂产卵策略时,须同时考虑寄主日龄的影响。
4.结论
本论文研究表明,除了寄主蛹体型大小外,寄主蛹日龄以及寄生蜂日龄也显著影响蝶蛹金小蜂的产卵决策。其他潜在因素(如抱卵量、身体营养状态、被捕食和寄生蜂风险等)对蝶蛹金小蜂产卵决策的影响仍有待于进一步研究。
Parasiotids are ideal model systems for testing behavioral and evolutionary hypotheses. Host choice is realized from interaction between host quality and host value in parasitoids. Host age and body size are the two main variables that constitute host quality, as manifested by fitness correlates of offspring parasitoids including clutch size, sex ratio, and body size. As of this time, the "host size-quality" hypothesis has been substantiated by numerous empirical studies of idiobiontic parasitoids. However, host age can influence the prediction of the hypothesis in idiobionts, as host quality may covary with host age. This thesis study, by using gregarious parasitoid Pteromalus puparum and host pupa of Pieris rapae as a model system, was intended to test the host size-quality hypothesis by manipulating host pupa size through starvation of larvae, host age, and parasitoid age as may affect oviposition decision in clutch size and sex ratio. The main results are as followed below.
1. Influence of host pupa weight on oviposition strategy of Pteromalus puparum
To test the "host size-quality"hypothesis, larvae of P. rapae were manipulated by starvation regime to obtain a wide range of body sizeofhost pupa, and then these puae were individually exposed to parasitism by the gregarious parasitoid.After a single attack the pupae were observed for emergence of offspring parasitoids. The results showed that the offspring production was linearly increased with increasing host pupa mass at oviposition, being 314.97 individuals per gram of pupa mass on average. The proportion of offspring females and the body size (hind tibia length) of offspring males and females also increased with increasing host pupa mass. The results suggest that P. puparum can allocate its offspring number and sex ratio, so as to maximize its fitness, basing on assessment of pupaquality. The reproductive strategy of P. puparum and related factors were discussed.
2. Effects of parasitoid age on oviposition strategy of Pteromalus puparum
To investigate how parasitoid age affects oviposition decision as predicted by the host size-quality hypothesis, parasitoids at either of 5-day,10-day,20-day, or 30-day old were provided with host pupae of different body sizes as obtained by the larval starvation. The results shown that parasitoid age significantly affect oviposition decision in P. puparum. With increase of parasitoid age, (ⅰ) clutch size obviously declined, with 49.14 (mean±se) from 5-day old mother wasps and 10.84from the 30-day old; (ⅱ) female increased in offspring, with 0.91 from 5-day old mother wasps and 0.95from the 30-day old; (ⅲ) body size enlarged slightly with 0.68mm in hind tibia length of females and 0.56mm of males from 5-day old mother wasps, and with 0.76mm of females and 0.61mm of males from 30-day old mother wasps. It is suggested that P. puparum might adjust oviposition decision in response to its own age.
3. Effects of host pupa age on oviposition strategy of Pteromalus puparum
To investigate effects of host pupa age on oviposition decision and offspring developmental performances of the parasitoid, host pupae at age of either 1-day,2-day, 3-day,4-day,5-day, and 6-day old were individually exposed to parasitism. The results shown that (ⅰ) clutch size decreased with host age; (2) female proportion declined with host age; (ⅲ) body size was not affected by host age from 1-day through to 5-day, but reduced from attacking 6-day old pupa with much reduced survival by 40%. It is suggested that host pupa age is an important variable in oviposition strategy of Pteromalus puparum, and therefore may confound predictions of the host-size-quality hypothesis.
4. Conclusions
The thesis study indicates that host pupa size is a main variable that measures host quality by gregarious Pteromalus puparum, but host age and mother wasp age are also important variables that affect host quality as perceived by the parasitoid as well. In addition to these variables, other potential traites, such as egg load, nutrition state, risks of predation and parasitism, etc may influence ovipisiton strategy, which are remained for further studies before fully understanding of oviposition strategy.
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38. Ellers J, van Alphen JJM, Sevenster JG.1998. A field study of size-fitness relationships in the parasitoid Asobara tabida. Journal of Animal Ecology,67:318-324.
39. Ellers J, van Alphen JJM.1997. Life history evolution in Asobara tabida:plasticity in allocation of fat reserves to survival and reproduction. Journal of Evolutionary Biology,10:771-785.
40. Ellers J, van Alphen JJM.2002. A trade-off between diapause duration and fitness in female parasitoids. Ecol.Entomol.27:84-279.
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44. Harvey JA, Bezemer TM, Elzinga JA, Strand MR.2004a. Development of the solitary endo-parasitoid Microplitis demolitor:host quality does not increase with host age and size. Ecological Entomology,29:35-43.
45. Harvey JA, Jervis MA, Gols GJZ, Jiang N, Vet LEM.1999. Development of the parasitoid:Cotesia rubecula (Hymenoptera:Braconidae) in Pieris rapae and P.brassicae (Lepidoptera:Pryalidae): evidence for host regulation. Journal of Insect Physiology,45:173-182.
46. Harvey JA, Strand MR.2002. The developmental strategies of endoparasitoid wasps vary with host feeding ecology. Ecology,83:2349-2451.
47. Harvey JA, van Dam NM, Gols R.2003. Interactions over four trophic levels:foodplant quality affects development of a hyperparasitoid as mediated through a herbivore and its primary parasitoid. Journal of Animal Ecology.72:529-531.
48. Harvey JA, Vet LEM, Jiang NQ, Gols R.1998. Nutritional ecology of the interaction between larvae of the gregarious ectoparasitoid, Muscidifurax raptorellus (Hymenoptera:Pteromalidae), and their pupal host, Musca domestica (Diptera:Muscidae). Physiological Entomology,23:113-120.
49. Harvey JA, Vet LEM, Witjes LMA, Bezemer TM.2005a. Remarkable similarity in body mass of a secondary hyperparasitoid Lysibia nana and its primary parasitoid host Cotesia glomerata emerging from cocoons of a comparable size. Archives of Insect Biochemistry and Physiology in press.
50. Harvey JA, Witjes LMA, Wagenaar R.2004b. Development of hyperparasitoid wasp Lysibia nana (Hymenoptera:Ichneumonidae) in a multitrophic framework. Environmental Entomology,33: 1488-1496.
51. Harvey JA, Strand MR.2002. The developmental strategies of endoparasitoid wasps vary with host feeding ecology. Ecology,83:2439-2451.
52. Heimpel GE, Rosenheim JA.1998. Egg limitation in parasitoids:a review of the evidence and a case study. Biol. Cont,11:160-168.
53. Honda JY, RF Luck,2001. Interactions between host attributes and wasp size:a laboratoryevalution of Trichogramma platneri as an augmentative biological control agent for two avocado pests. Entomologia Experimentalis et Applicata,100 (1):1-13.
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