玉米螟赤眼蜂自峰密度的干扰效应研究

玉米赤眼蜂自身密度干扰效应应显著,在同一寄主（米蛾卵）密度条件下,干扰效应符合Ｈａｓｓｅｌｌ模型,在不同寄主密度条件下,赤眼蜂的寄生功能反应符合Ｂｅｄｄｉｎｇｔｏｎ模型,自身密度对寄生率以及个体生殖力也存在于干扰效应。     

3种赤眼蜂对甜菜夜蛾和小菜蛾卵的寄生功能反应

为评估稻螟赤眼蜂Trichogramma japonicum Ashmead、螟黄赤眼蜂Trichogramma chilonis Ishii和玉米螟赤眼蜂Trichogramma ostriniae Pang et Chen对甜菜夜蛾Spodoptera exigua Hübner卵和小菜蛾Plutella xylostella Linnaeus卵的寄生潜能,本研究在室内条件下测试了3种赤眼蜂对不同密度甜菜夜蛾卵和小菜蛾卵的寄生率、寄生功能反应和搜寻效应。结果表明,稻螟赤眼蜂、螟黄赤眼蜂和玉米螟赤眼蜂对甜菜夜蛾卵和小菜蛾卵均有一定的寄生效应。螟黄赤眼蜂对甜菜夜蛾卵的寄生率、瞬间攻击率、寄生效能和搜寻效应均高于稻螟赤眼蜂和玉米螟赤眼蜂;3种赤眼蜂对小菜蛾卵的寄生率、瞬间攻击率、寄生效能和搜寻效应则表现为：稻螟赤眼蜂>螟黄赤眼蜂>玉米螟赤眼蜂。3种赤眼蜂对两种寄主的寄生功能反应均符合HollingⅡ型模型方程,表现为寄生量均随寄主卵密度的增加而升高,达到一定水平后趋于平稳。赤眼蜂的搜寻效应随寄主卵密度的增加而降低。综合分析可知,螟黄赤眼蜂对于甜菜夜蛾的控害能力优于稻螟赤眼蜂螟和玉米螟赤眼蜂,而稻螟赤眼蜂比螟黄赤眼蜂和玉米螟赤眼蜂更适合用于防控小菜蛾。     

长尾潜蝇茧蜂对橘小实蝇幼虫的寄生效能

以橘小实蝇幼虫为寄主,在(25±1)℃,RH(70±10)%,光照周期14L∶10D的条件下,研究了在不同密度的寄生蜂与寄主条件下的长尾潜蝇茧蜂寄生效能。结果表明:不同密度的橘小实蝇幼虫对长尾潜蝇茧蜂的寄生作用有明显的影响,其寄生效应符合功能反应模型HollingⅡ型,方程为Na=0.7291No/(1 0.02362No)。长尾潜蝇茧蜂自身密度对寄生作用也存在干扰效应,用Hassell-Varley模型可表示为a=0.1993P-0.2966。     

蝇蛹金小蜂对桔小实蝇蛹的寄生功能反应

在温度25±1℃、相对湿度70±5%、光照周期（L∶D）14 h∶10 h条件下,研究了蝇蛹金小蜂对桔小实蝇蛹的寄生功能反应及干扰效应。结果表明,寄主密度、寄主日龄均影响寄生蜂的寄生效能。蝇蛹金小蜂对桔小实蝇2日龄（N=2）、4日龄（N=4）蛹的寄生功能反应均符合HollingⅡ模型,其方程分别为Na= 0.4494N2/ (1+ 0.0294N2)、Na= 0.5586N4/ (1+ 0.0253N4)。24h内单头雌成蜂最多可寄生2日龄、4日龄蝇蛹数量分别为15.29、22.08头。自身密度对蝇蛹金小蜂寄生产生一定的干扰效应,其干扰效应符合Hassell－Varley模型（a= 0.0719P－0.2526）,表明蝇蛹金小蜂雌蜂的发现域随着自身密度的增加而逐渐变小,雌蜂个体间干扰效应降低了寄生效能。     

花角蚜小蜂对松突圆蚧的寄生功能反应

在实验室条件下,以松突圆蚧(Hemiberlesia pitysophilaTakagi)雌成蚧为寄主,研究了花角蚜小蜂(Coccobius azumaiTachikawa)的寄生功能反应.结果表明:寄生功能反应均符合HollingⅡ型方程,且功能反应受到温度、寄主密度和寄生物密度的影响.在同一温度下,寄生数量随寄主密度的增大而增加;在15℃~25℃范围内,随着温度的升高,被寄生的松突圆蚧雌成蚧数量增加,而在25℃~35℃之间呈相反趋势.花角蚜小蜂的寄生功能反应有较强的种内干扰作用,随自身密度的增加,寄生数量逐渐减少.Hassell(1969)模型E=QP-m和Bedding-ton(1975)模型E=aT/[1 btw(P-1)]均能较好地反映花角蚜小蜂的寻找效应与其自身密度之间的关系,模拟结果分别为E=0.1659P-0.5597和E=0.1437T/[1 0.2691(P-1)].     

长柄俑小蜂寄生橘小实蝇蛹的功能反应

长柄俑小蜂寄生橘小实蝇蛹的功能反应符合功能反应模型HollingⅡ型,其方程为:Na=1.5324TrN0/(1 1.5324ThN0)。每只寄生蜂在24 h内最大寄生量为17.57只橘小实蝇蛹。寄生1只寄主蛹所需的时间为1.37 h;瞬时攻击率(功能系数)为0.6526;长柄俑小蜂自身密度对寄生产生干扰效应,用Hassell-Varley模型可表示为:α=0.1085P-0.1929。     

蓝色长盾金小蜂对橡副珠蜡蚧的控制作用

蓝色长盾金小蜂Scutellista caerulea Fonscolombe是橡胶树重要害虫橡副珠蜡蚧Parasaissetia nigra Nietner的一种外寄生性天敌。为了明确蓝色长盾金小蜂对橡副珠蜡蚧的控害潜能,为该寄生蜂的进一步利用提供数据支撑。在室内通过体视镜下解剖观察寄生蜂在寄主腹下是否产卵测定了不同温度、寄主发育阶段下蓝色长盾金小蜂的寄生功能反应、寻找效应和自身密度干扰效应。温度设置有21、24、27、30、33、36℃共6个处理,寄主发育阶段设初期成虫（1-2 d成虫）、褐色期成虫（体色褐色,产卵前3-4 d成虫）、黑色期成虫（体色黑色,产卵1-2 d成虫）三个处理。结果显示：蓝色长盾金小蜂对橡副珠蜡蚧的寄生功能反应符合Holling-Ⅱ型和Holling-Ⅲ型模型,在21-36℃范围内,33℃时该蜂寄生效能最大,为44.4201,21℃时最小,为9.2458;在橡副珠蜡蚧为初期成虫-黑色期成虫范围内,寄生效能由大到小为黑色期成虫>褐色期成虫>初期成虫,分别为18.9044、13.7410、7.2002。采用Hassell-Varley干扰模型对不同温度下蓝色长盾金小蜂受自身密度干扰的寄生作用率进行拟合,表明蓝色长盾金小蜂在寄生时存在种群内个体间自我干扰情况。温度会影响蓝色长盾金小蜂的搜寻和自我干扰,在21-33℃范围内,33℃搜寻常数和干扰常数达到最大值,分别为0.6116、0.7535。当温度为33℃,寄主发育阶段为黑色期成虫时,蓝色长盾金小蜂对橡副珠蜡蚧有较强的控制能力,自身干扰作用最强。     

两种赤眼蜂对小菜蛾卵的寄生潜能分析

应用生命表技术分析了拟澳洲赤眼蜂Trichogramma confusum Viggiani(T.c)和卷蛾分索赤眼蜂Trichogramm-atoidea bactrae Nagaraja(T.b)在两种繁蜂条件组配下（分别用米蛾Corcyra cephalomica(Stainton)(RM)卵和小菜蛾Plutella xylostella(L.)(DBM)卵繁育）对小菜蛾卵的寄生潜能分析,结果表明：（1）在相同寄主繁蜂条件下,卷蛾分索赤眼蜂在小菜蛾卵上显示出较强的寄生潜能,米蛾卵上所繁的卷蛾分索赤眼蜂（T.b-RM)和小菜蛾卵上所繁的卷蛾分索赤眼蜂（T.b-DBM)的内禀增长率为0.3509和0.3450,而米蛾卵上所繁的澳洲赤眼蜂（T.c-RM)和小菜蛾卵上所繁的拟澳洲赤眼蜂（T.c-DBM)的仅为0.2391和0.1902,T.b-RM和T.b-DBM的每雌平均寄生卵数为70.75和46.13粒,而T.c-RM和T.c-DBM的仅为64.90和31.73粒,但拟澳洲赤眼蜂的雌蜂寿命较卷蛾分索赤眼蜂更长。（2）在不同寄主繁蜂条件下,同种赤眼蜂对小菜蛾卵的寄生潜能以米蛾卵所繁的仔蜂的各项寄生特性参数（内禀增长率,每雌寄生卵量,净生殖力,平均世代历期和雌蜂寿命）均优于用小菜蛾卵所繁之蜂,米蛾卵是其适宜的中间寄主。（3）长期用中间寄主繁蜂,赤眼蜂对目标寄主表现出一定的不适应性,中间寄主的驯化对赤眼蜂的寄生潜能有不容忽视的削弱作用。     

赤眼蜂寄主偏爱可塑性研究

三种赤眼蜂寄主选择性和寄生能力的实验结果表明,松毛虫赤眼蜂对亚洲玉米螟卵的寄主选择性和寄生能力最弱,其次为拟澳洲赤眼蜂,玉米螟赤眼蜂最强.寄主偏爱可塑性实验结果表明,对玉米螟卵选择性和寄生能力较弱的拟澳洲赤眼蜂,其寄主的低偏爱性是可塑的;而对玉米螟卵选择性和寄生能力较强的玉米螟赤眼蜂,在亚洲玉米螟卵繁殖几代后,其寄生能力变化不大.     

繁育寄主对稻螟赤眼蜂寄生行为及寄生能力的影响

【目的】繁育寄主影响赤眼蜂的生物学特性,为明确米蛾 Corcyra cephalonica (Stainton)卵繁育的稻螟赤眼蜂 Trichogramma japonicum Ashmead是否和二化螟Chilo suppressalis (Walker)卵繁育稻螟赤眼蜂在防治二化螟上有差别,及田间用米蛾卵作为稻螟赤眼蜂续代寄主的可行性,本实验观察了羽化自不同寄主的稻螟赤眼蜂的寄主选择和寄生行为,研究了两种繁育寄主对稻螟赤眼蜂寄生能力的影响,为田间防治水稻螟虫提供参考信息。【方法】室内利用选择试验观察了由米蛾卵和二化螟卵繁育出的稻螟赤眼蜂对两种寄主的选择趋性及在两种寄主上的寄生行为和寄生能力,统计了羽化子代蜂的寿命。【结果】由二化螟卵繁育的稻螟赤眼蜂表现出了对二化螟卵的选择偏好,而米蛾卵繁育的稻螟赤眼蜂在米蛾卵和二化螟卵间没有表现出寄主选择偏好。寄主搜寻时间不受繁育寄主及供试寄主的影响,卵表探测时间和穿刺及产卵时间不受繁育寄主的影响,但同一寄主繁育的稻螟赤眼蜂在米蛾卵上的卵表探测时间显著长于在二化螟卵上的卵表检测时间。除二化螟卵繁育蜂寄生二化螟卵时的寄生率（36.95%）和羽化率（45.68%）较低外,米蛾卵繁育蜂寄生米蛾卵或二化螟卵以及二化螟卵繁育蜂寄生米蛾卵等3组处理间的寄生率、羽化率均无显著差异,各处理组间子代蜂寿命也无显著差异。【结论】繁育寄主影响稻螟赤眼蜂的寄主选择和寄生能力,但不影响寄生蜂搜寻寄主的能力和寄生行为。     

On the capacity of macroparasites to control insect populations

A graphical model of the population dynamics of macroparasites and their hosts is developed. Three principal means by which the parasites can be regulated are considered: reduction in host density as a result of parasite-induced host mortality, reduction in host density as a result of parasite-induced host sterility, and competition among parasites within multiply-infected hosts. The means by which parasites are regulated has a major effect on the degree to which they can depress host population densities. In particular, a parasite that sterilizes its host is expected to reduce host density more than one that causes an equivalent decline in host fitness through increased mortality. A special case of the model is developed for herbivorous insects that, in the absence of parasites, are limited by larval food resources. Parasites that are regulated via parasite-induced host sterility will control the insect populations below the level set by larval resources if the threshold host density for the parasites (N(T)) is less than the ratio of carrying capacity to net reproductive rate of the insects (K/R). Data are presented showing that all three means of parasite regulation, but especially parasite-induced host sterility, can operate in Howardula aoronymphium, a nematode parasite of mycophagous Drosophila flies. Data from a field cage experiment show that, if these nematodes are regulated primarily via reductions in host density due to this sterility, the parameters N(T), K, and R are such that Howardula is likely to play an important role in controlling Drosophila populations. However, this conclusion must be tempered by the fact that these nematodes also cause increased host mortality and experience within-host competition, making the conditions for parasite control of the flies more stringent.     

Intraspecific competition and density dependence in an Ephestia kuehniella–Venturia canescens laboratory system

A model host-parasitoid system of Ephestia kuehniella and Venturia canescens was used to examine the influence of host and parasitoid density on host and parasitoid life-history parameters via a two-way factorial experimental design (5 initial host densities×3 parasitoid densities). In the absence of parasitoids, E. kuehniella experienced scramble-type competition with reduced growth, diminished adult size and a subsequent fecundity trade-off for mortality. The mortality that did occur was confined to the late larval and pupal stages. In the presence of parasitoids attacking the late larval stage, competition changed from scramble for food to contest for enemy-free space, with hosts escaping parasitism being small with low fecundity and reduced egg size, and with parasitoid adult size inversely dependent on host density. Total insect emergence (host+parasitoid), a measure of the influence of host resource competition on survivorship, exhibited a threshold effect as a function of initial host density; the threshold value was increased to a higher initial host density in the presence of parasitoids. Models of host self-limitation were fitted to the data, with the generalized Beverton-Holt model that incorporates a threshold effect providing the best fit, and the Ricker model with no threshold providing a very poor fit to the data.     

Reproduction now or later: optimal host-handling strategies in the whitefly parasitoid Encarsia formosa

We developed a dynamic state variable model for studying optimal host‐handling strategies in the whitefly parasitoid Encarsia formosa Gahan (Hymenoptera: Aphelinidae). We assumed that (a) the function of host feeding is to gain nutrients that can be matured into eggs, (b) oögenesis is continuous and egg load dependent, (c) parasitoid survival is exponentially distributed and (d) parasitoids encounter hosts randomly, are autogenous and have unlimited access to non‐host food sources to obtain energy for maintenance and activity. The most important prediction of the model is that host feeding is maladaptive under field conditions of low host density (0.015 cm^?2) and short parasitoid life expectancy (maximum reproductive period of 7 d). Nutrients from the immature stage that can be matured into eggs are sufficient to prevent egg limitation. Both host density and parasitoid life expectancy have a positive effect on the optimal host‐feeding ratio. Parasitoids that make random decisions gain on average only 35% (0.015 hosts cm^?2) to 60% (1.5 hosts cm^?2) of the lifetime reproductive success of parasitoids that make optimal decisions, independent of their life expectancy. Parameters that have a large impact on lifetime reproductive success and therefore drive natural selection are parasitoid life expectancy and the survival probability of deposited eggs (independent of host density), the number of host encounters per day (when host density is low) and the egg maturation rate and number of host types (when host density is high). Explaining the evolution of host‐feeding behaviour under field conditions requires field data showing that life expectancy in the field is not as short as we assumed, or may require incorporation of variation in host density. Incorporating variation in walking speed, parasitised host types or egg resorption is not expected to provide an explanation for the evolution of host‐feeding behaviour under field conditions.     

The impact of resource limitation and the phenology of parasitoid attack on the duration of insect herbivore outbreaks

Fluctuations in resource quality and quantity, and changes in mortality due to predators and parasites are thought to be of prime importance in the regular fluctuations of forest insects. We examine how food limitation and parasitoids with different phenologies of attack regulate the population cycles of insect hosts. Our analysis of the limit cycle of a model with a biologically realistic form of density dependence in the host yields two novel predictions. First, outbreaks will typically last for only 2 generations after parasitoids begin to reduce the host population below the maximum density. Second, host growth rate is important in determining cycle length only when parasitoids attack before the impacts of resource limitation affect the host. The robustness of these predictions are tested using a more general form of density dependence in the host, revealing that our predictions are valid as long as density dependence in the host is not too overcompensatory.     

A simple model of host-parasitoid interaction with host-feeding

Summary A simple mathematical model of host-parasitoid interaction with host-feeding was presented with special reference to the system of the greenhouse whitefly and the parasitoidEncarsia formosa. In the model, when a parasitoid encounters a host, it has a choice between feeding the host and ovipositing one egg in the host. It was shown that an intermediate value of the feeding ratio of all attacks gives the minimum equilibrium host density and the minimum amplitudes of fluctuation in the densities of the two species. Computer simulations of a modified model with time lags also gave the similar results. The model suggested for natural enemy introduction program that parasitoid species with host-feeding habits are promising agents for effective controls for pest insects and that the timing of introduction is very important. By an evolutionary analysis, it was shown that the feeding ratio evolves to minimize the host density under natural selection among parasitoids.     

Parasitoid aggregation and interference in host–parasitoid dynamics

1. Parasitoids do not distribute themselves evenly among available patches, which has an important bearing on the dynamics of host–parasitoid interactions. This study examined the density‐dependent nature of aggregation of the parasitoid Dirhinus giffardii Silvestri on the oriental fruit fly host, Bactrocera dorsalis (Hendel) distributed among discrete patches. 2. Four artificial patches were created in a cage, and the number of hosts in each patch was manipulated. Parasitoids were released into the cage, and whether parasitoid density and host density influence the degree of parasitoid aggregation was examined. 3. Parasitoid aggregation became stronger (e.g. uneven distribution among patches) as the parasitoid density decreased and also as the host density increased. The index of parasitoid aggregation was not influenced by the distribution of hosts among patches. 4. The empirically characterised aggregation pattern was incorporated in a host–parasitoid model that consists of one host and one parasitoid species. The analysis of the model shows that an unstable system (i.e. the coexistence of the host and parasitoid is impossible) can be stabilised (i.e. coexistence is possible) when the parasitoid aggregates in a way that is consistent with the pattern found in the experiment.     

Evolution of airborne infectious diseases according to changes in characteristics of the host population

The authors predicted evolutionary changes in airborne infectious diseases according to changes in the characteristics of the host population. The predictions were based upon a mathematical model of infectious diseases and the validity of the predictions was verified against the history of man and pathogens. The feature of this model is that it involves a density of pathogens in the environment as an additional variable which can be regarded as more suitable to airborne infectious diseases. In spite of this modification, this study reached a similar conclusion to the threshold density theory: that is, susceptible host density in the absence of the pathogen must be larger than that in the presence of the pathogen, for the pathogen to be persistent. Moreover the authors concluded that one type of pathogen cannot be replaced by another type of pathogen as long as the susceptible host density of the former type is the mininum one. The predictions were considered to be valid for a wide range of infectous diseases. Making use of these principles, the authors predicted that the variety of infectious diseases should increase as host density increases and that pathogens should evolve to be less virulent as the host life-span increases. The finalidea discussed is whether or nor the history of man and pathogen can be verified by the predictions.     

Egg Distributions of Insect Parasitoids: Modelling and Analysis of Temporal Data with Host Density Dependence

A simple numerical procedure is presented for the problem of estimating the parameters of models for the distribution of eggs oviposited in a host. The modelling is extended to incorporate both host density and time dependence to produce a remarkably parsimonious structure with only seven parameters to describe a data set of over 3,000 observations. This is further refined using a mixed model to accommodate several large outliers. Both models show that the level of superparasitism declines with increasing host density, and the rate declines over time. It is proposed that the differing behaviours represented by the mixed model may reflect a balance between behavioural strategies of different selective benefit.     

Populations and infectious diseases: Dynamics and evolution

The host-parasite or host-pathogen system was analyzed from dynamical and evolutionary viewpoints using simple mathematical models incorporating vertical transmission, immunity and its loss. We first analyzed a model without density regulation of host population. In the analysis on dynamics, the condition for the pathogen to work as a density regulating factor was obtained. In the analysis on evolution, criteria for the evolution of host and pathogen were proposed. These criteria implies that the evolution of hosts should result in an increase in infected host density, whereas the evolution of pathogens a decrease in susceptible host density. The direction of evolution at some parameters of host and that of pathogen were examined when the parameters were independently and freely changeable. Among the parameters, only reduction in additional mortality due to infection was the evolutionary trend common to both host and pathogen. In all the other parameters examined, trend of evolution predicted in host is reversed in pathogen. We then analyzed whether the obtained criteria still hold in models with density regulation of hosts. Using randomly generated parameter sets, we obtained the result that the criteria should hold very likely though they do not always hold. We discussed evolution of virulence when there is a constraint between the traits.     

Optimal foraging predicts the ecology but not the evolution of host specialization in bacteriophages

We explore the ability of optimal foraging theory to explain the observation among marine bacteriophages that host range appears to be negatively correlated with host abundance in the local marine environment. We modified Charnov's classic diet composition model to describe the ecological dynamics of the related generalist and specialist bacteriophages phiX174 and G4, and confirmed that specialist phages are ecologically favored only at high host densities. Our modified model accurately predicted the ecological dynamics of phage populations in laboratory microcosms, but had only limited success predicting evolutionary dynamics. We monitored evolution of attachment rate, the phenotype that governs diet breadth, in phage populations adapting to both low and high host density microcosms. Although generalist phiX174 populations evolved even broader diets at low host density, they did not show a tendency to evolve the predicted specialist foraging strategy at high host density. Similarly, specialist G4 populations were unable to evolve the predicted generalist foraging strategy at low host density. These results demonstrate that optimal foraging models developed to explain the behaviorally determined diets of predators may have only limited success predicting the genetically determined diets of bacteriophage, and that optimal foraging probably plays a smaller role than genetic constraints in the evolution of host specialization in bacteriophages.