捕食风险对越冬根田鼠肠道寄生物易感性的影响

在自然生态系统中,不同营养级物种可通过特征介导间接效应对生态系统的稳定及种群产生深刻的影响。但目前有关特征介导间接效应的实验研究多见于无脊椎动物、鱼类和两爬类。本研究以根田鼠为对象,在野外围栏内建立预防捕食者和未预防捕食者两种实验处理种群,并通过采用麦克马斯特法测定两种处理种群实验个体肠道寄生物感染种类及感染率和感染强度,采用PHA（phytohemagglutinin）反应和白细胞分类计数测定不同处理种群实验个体免疫能力,以分析捕食风险对根田鼠肠道寄生物的感染效应。结果表明,未预防捕食者处理组根田鼠PHA反应、白细胞计数和淋巴细胞计数较预防捕食者处理组实验个体显著降低,而球虫 E. wenrichi 的感染率和感染强度则显著增加,但绦虫和线虫以及其他3种球虫的感染率和感染强度无显著差异。结果表明,捕食者可通过介导猎物免疫力特征而间接影响猎物肠道寄生物的感染,验证了本项提出的捕食风险可通过降低根田鼠的免疫能力而增加其肠道寄生物感染的假设。     

野外围栏条件下根田鼠肠道寄生物的感染率

寄生物是调节小哺乳动物种群波动的因子之一。许多生物和非生物因子可影响寄生物的感染。本研究以根田鼠为对象,在野外围栏条件下测定了根田鼠肠道寄生物种类及其感染率,并分析了根田鼠建群者数量、性别、年龄及时间对寄生物感染率的影响。结果表明,根田鼠肠道寄生物有球虫、线虫和绦虫,球虫是主要寄生物, Eimeria wenrichi 为球虫中的优势种;其感染率随根田鼠建群者数量增加而增加,且随时间而显著变化。在根田鼠幼体中,高密度处理组E. wenrichi 的感染率显著高于低密度处理组,而根田鼠幼体的E. ochrogasteri 感染率显著高于成体。本研究未发现根田鼠性别间的球虫感染率差异。上述结果说明,在根田鼠种群中,球虫感染率有时间变化,且受宿主密度的制约,高密度种群引起的生前应激与当前密度制约性感染可能对球虫感染率具有耦合效应。     

艾美耳球虫对高原鼠兔繁殖的影响

寄生物对宿主繁殖的影响取决于宿主对当前繁殖值和剩余繁殖值的权衡。球虫为微型寄生物,而微型寄生物对宿主当前繁殖值的影响较剩余繁殖值要大。因此,本研究检验了寄生在高原鼠兔肠道内的艾美耳球虫可影响其当前繁殖的假设。在繁殖早、中、晚期,野外共观测高原鼠兔170只。结果表明,不同繁殖期感染率有显著差异。在繁殖中期,未感染雌性的妊娠率显著高于感染雌性。且未妊娠雌性较妊娠雌性有更高的感染强度,但在另外两个繁殖期没有发现此效应。在雄性中,任何繁殖期的感染强度和感染率与睾丸和附睾指数均无显著相关性,且感染和未感染球虫雄性睾丸及附睾指数无显著差异。此外,野外观测实验结果表明,感染雌性的胚胎重较未感染雌性显著降低,与野外感染对胚胎重量影响的实验结果相一致。说明艾美耳球虫感染可影响胚胎的发育。上述结果说明,艾美耳球虫对高原鼠兔繁殖的影响随繁殖期而有不同效应,且存在性别间差异,这种效应可能与不同性别间的繁殖对策有关。     

食物、捕食和种间竞争对东方田鼠种群动态的作用

采用2×2×2析因实验设计,在野外围栏条件下,测定食物、捕食和竞争物种黑线姬鼠(Apodemus agrarius)对东方田鼠(Microtus fortis)种群动态作用的格局.食物可利用性、捕食及种间竞争的独立作用对种群最小存活数均具有极显著的效应,除捕食与种间竞争的交互作用接近显著水平外,食物与种间竞争、食物与捕食者以及三者间的交互作用均不显著;三类外部因子对种群补充量的独立作用效应均达到极显著水平,且对种群补充量的作用具有累加效应;食物可利用性、捕食及种间竞争对种群繁殖成体的比例具有极显著的作用;三类外部因子对种群幼体与成体的比例具有极显著的作用.对种群年龄结构而言,与捕食者及种间竞争比较,食物可利用性是相对较弱的影响因子,在任何捕食与种间竞争交互作用条件下,食物的作用均不显著;三类外部因子均能显著地影响东方田鼠的体重增长率,但三者的交互作用对其影响不显著;MANOVA结果表明,捕食对成体存活率的作用最强烈,其次,为食物可利用性,种间竞争的作用最弱,但三者的交互作用效应不显著.对幼体的存活时间,除捕食的作用接近显著水平外,食物可利用性及种间竞争的作用均不显著.结果提供了食物可利用性、捕食和种间竞争对东方田鼠种群动态作用的充分证据,验证了食物、捕食和种间竞争对田鼠类种群动态具有独立或累加效应的总假设.     

根田鼠攻击行为模式及其进化稳定对策分析

在捕食和附加食物交互作用条件下 ,测定根田鼠种群攻击水平与种群参数之间的关系 ,应用对策论分析攻击行为的模式。通过观测 1 6 5对实验个体的攻击行为 ,发现体重和繁殖特征决定根田鼠个体的攻击水平。体重较大个体及繁殖个体的攻击频次最高。攻击水平的性别差异显著。Kruskal- Wallis单侧秩方差检验结果表明 ,雄体攻击水平显著高于雌体。攻击水平与种群密度呈负相关关系 ,但不显著。与留居个体比较 ,扩散个体的攻击性弱。高质量的附加食物可降低根田鼠的攻击水平 ,捕食者的存在不直接影响攻击水平。根田鼠的攻击行为属仪式化非伤害行为。ESS的鹰 -鸽对策模型分析结果与实验结果一致 :不同处理种群 ESS集的鹰型对策者比例有显著差异 ,环境条件愈差 ,取胜的价值愈大 ,种群中鹰型对策者比例愈高。有捕食者、无附加食物( P,- F)种群的环境条件最差 ,鹰型对策者比例最高 ,为 0 .4 0 ;次为无捕食者、无附加食物 (- P,- F)种群 ,为 0 .33;有捕食者、有附加食物 ( P, F)种群 ,为 0 .1 8;无捕食者、有附加食物 (- P, F)种群的环境条件最优越 ,鹰型对策者比例最低 ,为0 .1 7。验证了捕食和食物交互效应能改变田鼠类动物攻击行为格局及对策模式的假设     

母体密度应激与球虫感染对子代根田鼠免疫力的影响

免疫功能是影响动物个体适合度的重要因素之一,也是防御病原体入侵的重要途径,对动物的生存至关重要。种群密度与寄生物感染都能影响动物的免疫功能。本实验通过2×2析因实验设计,测定了母体密度应激+球虫感染、母体密度应激+未球虫感染、未母体密度应激+球虫感染、未母体密度应激+未球虫感染等4个处理对根田鼠粪便皮质酮、球虫感染率、白细胞总数、各型白细胞的百分比、淋巴细胞亚型计数（CD4,CD8计数）等指标。结果表明,高密度母体应激处理可显著降低CD4数量、CD4/CD8比例、淋巴细胞数、嗜酸性粒细胞数、白细胞数,母体应激与球虫的耦合可使上述免疫指标进一步降低,说明母体密度应激和球虫感染对于根田鼠的免疫具有负的叠加效应。     

小型啮齿动物种群系统调节复合因子理论的野外实验研究：食物可利用性和捕食对根田鼠种群动态作用的分析

本研究在野外围栏条件下采用析因实验设计,测定营养、捕食及空间行为对根田鼠（Ｍｉ－ｃｒｏｔｕｓｏｅｃｏｎｏｍｕｓ）种群统计特征的影响。本文旨在检验下述特定假设：高质量食物可利用性和捕食对限制小型啮齿动物种群密度具有独立的和累加的效应。３年期间,４种野外实验处理６个重复的研究结果表明,附加食物并预防捕食者处理的种群具有最高密度;未附加食物及不预防捕食者处理（对照）的种群密度最低;而单一处理的种群,其密度居中。不同处理条件下,新生个体在种群的补充模式以及种群瞬时增长率的变化均与种群密度的变动相应一致。双因素ＡＮＯＶＡ的结果证明,附加高质量食物能明显地提高根田鼠的种群密度,而对种群补充量的作用则较弱,仅接近显著水平;预防捕食者不仅能显著地作用于种群密度,更能强烈地影响种群补充量。高质量食物和捕食者的作用具有累加的性质,两者的交互作用对种群密度和补充量均无显著影响。     

小型啮齿动物种群系统调节复合因子理论的野外实验研究：食物可利用性和捕食对根田鼠种群空间行为的作用模式及其对种群调节的探讨

本项研究在野外围栏条件下,采用析因实验设计,测定食物可利用性和捕食对根田鼠（Ｍｉｃｒｏｔｕｓｏｅｃｏｎｏｍｕｓ）种群空间行为的作用模式。检验的特定假设为,高质量食物较大的可利用性能降低田鼠的攻击行为和活动;捕食能减少田鼠的活动。研究结果表明,食物可利用性能间接地和直接地影响根田鼠的空间行为。附加食物种群具有较高的密度和较小的巢区,且在诱捕期间具有较少的长距离活动和较低的攻击水平。捕食者的存在不直接影响攻击行为,但能影响诱捕期间的长距离活动,此为根田鼠对捕食者存在作出的直接反应。在阐明田鼠种群动态时,应仔细考虑上述因子相互作用的效应。     

田鼠种群波动的原因和时间

本文总结了橙腹田鼠（Microtus ochrogaster）和草原田鼠（M．permsylvanicus）25年的种群统计学研究结果和结论。探讨了田鼠种群波动周期性、诱发种群波动以及导致波动期间峰值变异的因素。并对种群存活值和繁殖活动的作用进行了分析和评价。根据两种田鼠种群波动周期性、波动峰值出现的时间以及特定年份峰值的高度等特征,证明两物种波动均具有不稳定性。两种田鼠存活值的变化是由特定年份是否发生波动以及波动峰值出现的时间决定。增加初始阶段的种群密度及时间长度是造成两种动物种群波动峰值不同的主要原因。橙腹田鼠种群停止增长的原因是存活值降低,而草原田鼠则是繁殖活动减少。据推测,与种群波动初始密度相关的种群死亡率的差异是由捕食者的净效应（Net effect）决定的,调控两种群密度的因素均为非密度的其它生态学因子。由于特定年份田鼠种群捕食压力的不确定性,导致了橙腹田鼠和草原田鼠种群波动的不稳定性。     

捕食和食物交互作用条件下根田鼠季节性波动种群攻击水平及其行为多态性分析

在捕食和附加食物交互作用条件下,以恐吓、进攻、追逐、争斗及回避5类行为为变量,以恐吓、进攻、追逐计数之和的平均值作为攻击水平,测定根田鼠种群不同波动时期成体的攻击水平.发现根田鼠的攻击性与种群波动时期之间,存在明显的关联.统计分析结果表明,在种群3个波动时期,4种处理种群两性攻击型个体比例差异显著.除预防捕食者无附加食物（-P,-F）种群的雌体外,其它处理种群增长期和高峰期雌性和雄性攻击型个体的比例高于其衰减期.其中,预防捕食者附加食物（-P,＋F）种群、-P,-F种群及未预防捕食者附加食物（＋P,＋F）种群,雄性攻击型个体的比例均为增长期＞高峰期＞衰减期;在未预防捕食者无附加食物（＋P,-F）种群,雄性攻击型个体比例为高峰期＞增长期＞衰减期.各处理种群雌性攻击型个体比例的格局与雄体的不同.其中,-P,＋F种群及＋P,＋F种群为增长期＞高峰期＞衰减期,＋P,-F种群为高峰期＞增长期＞衰减期,而-P,-F种群攻击型个体比例为高峰期＞衰减期＞增长期.虽不同处理种群雌体及雄体的5类行为变量与种群密度的相关性不一致,而具有明显攻击性的恐吓、进攻及争斗3类行为则分别与种群密度呈显著或极显著的线性正相关关系,其结果与Chitty多态行为假设预测的一致;验证了所提出的特定假设：种群外部因子捕食和食物交互效应介导的攻击行为选择,是引起田鼠类种群季节性波动的主要内部因子.     

Parasite-induced parthenogenesis in a freshwater snail: stable,persistent patterns of parasitism

Summary The role of parasites in the evolution of host reproductive modes has gained renewed interest in evolutionary ecology. It was previously argued that obligate parthenogenesis (all-female reproduction) arose in a freshwater snail, Campeloma decisum, as a consequence of severe sperm limitation caused by an unencysted trematode, Leucochloridiomorpha constantiae. In the present study, certain conditions are examined for parasitic castration to account for the maintenance of parthenogenesis: the spatial patterns of the prevalence and intensity of infection on a broad geographical scale and its relationship to host genotype; the recovery from infection after isolation from sources of infection; age-related patterns of infections; and the effects of L. constantiae on snail fecundity.In contrast to the common pattern of the aggregated distribution of parasites within host populations, many snail populations with high prevalence and intensity of infection have non-aggregated parasite distributions. Clonal genotype of the host explained little of the variation in intensity and prevalence of infection by the parasite. Female snails maintained similar prevalence and intensity of infection after isolation, and individuals accumulated parasites throughout their lifespan, both of which suggest there is no effective immune response to infection by L. constantiae. Snail fecundity is not significantly influenced by the intensity of infection. These results suggest that L. constantiae may have represented a strong selective force against males during the initial introduction of this parasite into sexual snail populations because of the persistent nature of infection.     

野外条件下球虫感染对根田鼠免疫功能的影响

动物的免疫功能是影响动物适合度的重要因素之一。本文以根田鼠（Microtus oeconomus）为研究对象,在野外围栏条件下,通过球虫感染和混合驱虫药处理,测定根田鼠的植物血凝素反应（Phytohemagglutinin,PHA）、血清抗匙孔血蓝蛋白（Keyhole limpet haemoeyanin,KLH）的IgG 水平和血象参数,探讨寄生物感染对免疫功能的影响。结果表明,根田鼠随球虫感染率和感染强度的增加,对PHA的反应峰值降低、血清抗KLH的IgG水平、白细胞总数、淋巴细胞的百分比及血浆蛋白含量明显下降、嗜碱性粒细胞的百分比增加,嗜酸性粒细胞、嗜中性粒细胞和单核细胞的百分比未检出显著变化。由此说明,球虫感染降低了根田鼠的细胞免疫和体液免疫功能。     

A cost of Wolbachia-induced sex reversal and female-biased sex ratios: decrease in female fertility after sperm depletion in a terrestrial isopod

A number of parasites are vertically transmitted to new host generations via female eggs. In such cases, host reproduction is an intimate component of parasite fitness and no cost of the infection on host reproduction is expected to evolve. A number of these parasites distort host sex ratios towards females, thereby increasing either parasite fitness or the proportion of the host that transmit the parasite. In terrestrial isopods (woodlice), Wolbachia bacteria are responsible for sex reversion and female-biased sex ratios, changing genetic males into functional neo-females. Although sex ratio distortion is a powerful means for parasites to increase in frequency in host populations, it also has potential consequences on host biology, which may, in turn, have consequences for parasite prevalence. We used the woodlouse Armadillidium vulgare to test whether the interaction between Wolbachia infection and the resulting excess of females would limit female fertility through the reduction in sperm number that they receive from males. We showed that multiple male mating induces sperm depletion, and that this sperm depletion affects fertility only in infected females. This decrease in fertility, associated with male mate choice, may limit the spread of Wolbachia infections in host populations.     

Bot fly parasitism of the red-backed vole: host survival,infection risk,and population growth

Parasites can play an important role in the dynamics of host populations, but empirical evidence remains sparse. We investigated the role of bot fly (Cuterebra spp.) parasitism in red-backed voles (Myodes gapperi) by first assessing the impacts of the parasite on the probability of vole survival under stressful conditions as well as on the reproductive activity of females. We then identified the main factors driving both the individual risk of infection and the abundance of bot flies inside red-backed voles. Finally, we evaluated the impacts of bot fly prevalence on the growth rate of vole populations between mid-July and mid-August. Thirty-six populations of red-backed voles were sampled in the boreal forest of Québec, Canada. The presence and the abundance of parasites in voles, two host life history traits (sex and body condition), three indices of habitat complexity (tree basal area, sapling basal area, coarse woody debris volume), and vole abundance were considered in models evaluating the effects of bot flies on host populations. We found that the probability of survival of red-backed voles in live traps decreased with bot fly infection. Both the individual risk of infection and the abundance of bot flies in red-backed voles were driven mainly by vole abundance rather than by the two host life history traits or the three variables of habitat complexity. Parasitism had population consequences: bot fly prevalence was linked to a decrease in short-term growth rate of vole populations over the summer. We found that bot flies have the potential to reduce survival of red-backed voles, an effect that may apply to large portions of populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Regulation of root vole population dynamics by food supply and predation: a two-factor experiment

This paper reports the effects of food supply, predation and the interaction between them on the population dynamics of root voles, Microtus oeconomus , by adopting factorial experiments in field enclosures. This two-factor experiment proved the general hypothesis that food supply and predation had independent and additive effects on population dynamics of root voles. The experimental results proved the following predictions: (1) predation reduced population density and recruitment significantly; (2) food supply increased population density; (3) predation and food supply influenced spacing behavior of root voles separately and additively: Exposure to predation reduced long movements of root voles between trapping sessions; additional food supply reduced aggression level and home range size of root voles. Less movement of individuals that exposed to predators possibly reduced their opportunity of obtaining food and lessened population survival rate, which led population density to decrease. Smaller home range and lower aggression level could make higher population density tolerable. The interactive effect of predation and food on home range size was highly significant (P=0.0082<0.01). The interactive effect of food and predation on dispersal rate was significant (P<0.01). From the experimental results, we conclude that the external factors (predation, food supply) were more effective than internal factors (spacing behavior) in determining population density of root voles – under the most favorable external conditions (−P, +F treatment), the mean density and mean recruitment of root vole population was the highest; under the most unfavorable external conditions (+P, −F treatment), the mean density and mean recruitment of root vole population was the lowest.     

Patterns of parasite abundance and distribution in island populations of Galápagos endemic birds

Parasite life-history characteristics, the environment, and host defenses determine variation in parasite population parameters across space and time. Parasite abundance and distribution have received little attention despite their pervasive effects on host populations and community dynamics. We used analyses of variance to estimate the variability of intensity, prevalence, and abundance of 4 species of lice (Insecta: Phthiraptera) infecting Galápagos doves and Galápagos hawks and 1 haemosporidian parasite (Haemosporida: Haemoproteidae) infecting the doves across island populations throughout their entire geographic ranges. Population parameters of parasites with direct life cycles varied less within than among parasite species, and intensity and abundance did not differ significantly across islands. Prevalence explained a proportion of the variance (34%), similar to infection intensity (33%) and parasite abundance (37%). We detected a strong parasite species-by-island interaction, suggesting that parasite population dynamics is independent among islands. Prevalence (up to 100%) and infection intensity (parasitemias up to 12.7%) of Haemoproteus sp. parasites varied little across island populations.     

Effects of intensity and duration of infection by a hemiparasitic plant, Rhinanthus serotinus, on growth and reproduction of a perennial grass, Agrostis capillaris

Arising from annual variation in parasitic plant population densities, substantial yearly changes may occur in the parasitic load of an individual perennial host. We conducted two two-year greenhouse pot experiments to examine the effects of varying intensities and duration of infection by an annual root hemiparasitic plant. Rhinanthus serotinus, on the growth and reproduction of its perennial host grass. Agrostis capillaris. In the first experiment, one host plant was growing either alone or under a load of 1 or 3 root hemiparasitic plants for one growing season, and during the next season all hosts continued their life free of hemiparasites. In the second experiment, the host plants either grew alone or were parasitised by 1 or 2 root hemiparasitic plants either during the first growing season only or during two successive seasons (the parasitic load being the same in the two seasons). In both experiments, the root hemiparasites markedly reduced the growth and reproduction of their perennial hosts. In the first experiment, the negative effects of parasites on host performance increased with the increase in intensity of parasitic infection from one to three parasites. The harmful effects of hemiparasitim were carried over to the following season; hosts parasitised during the previous season with one or three parasites produced significantly less biomass than those without parasites. In addition, hosts parasitised by three parasites during the first season produced significantly less panicles in the second season than those parasitised by one parasite and those without parasites. The second experiment showed that the production of biomass of A. capillaris during the second season was, but the production of panicles was not affected by the duration of parasitic infection. In addition, in this experiment, the second season biomass of A. capillaris depended on the intensity of infection (1 vs 2 parasites), but the production of panicles was unaffected by the number of parasites.