Haploid and diploid survival differences demonstrate selection in scale insect demes

Summary Black pineleaf scale insect populations are subdivided into genetically differentiated demes associated with individual pine trees. A comparison of sex ratios early and late in the life cycle demonstrated differences in the mortality experienced by haploid males and diploid females. Hatching ratios were significantly female-biased, and differential mortality increased this bias in ratios estimated just before adult male eclosion. The relative survival of males and females varied with overall mortality, causing a correlation between local densities and the surviving sex ratio. We suggest (a) that the genetic differentiation of scale demes results in part from selection pressures associated with individual pine trees, (b) that this differentiation entails an accumulation of locally adaptive traits within the scale subpopulation on each tree, (c) that expression of these adaptations in the haploid and diploid sexes may vary with their frequencies, and (d) that the surviving sex ratio thus offers a comparative measure of selection and the local adaptation achieved by the insects in individual demes.     

The Effects of Background and Interference Selection on Patterns of Genetic Variation in Subdivided Populations

It is well known that most new mutations that affect fitness exert deleterious effects and that natural populations are often composed of subpopulations (demes) connected by gene flow. To gain a better understanding of the joint effects of purifying selection and population structure, we focus on a scenario where an ancestral population splits into multiple demes and study neutral diversity patterns in regions linked to selected sites. In the background selection regime of strong selection, we first derive analytic equations for pairwise coalescent times and F_ST as a function of time after the ancestral population splits into two demes and then construct a flexible coalescent simulator that can generate samples under complex models such as those involving multiple demes or nonconservative migration. We have carried out extensive forward simulations to show that the new methods can accurately predict diversity patterns both in the nonequilibrium phase following the split of the ancestral population and in the equilibrium between mutation, migration, drift, and selection. In the interference selection regime of many tightly linked selected sites, forward simulations provide evidence that neutral diversity patterns obtained from both the nonequilibrium and equilibrium phases may be virtually indistinguishable for models that have identical variance in fitness, but are nonetheless different with respect to the number of selected sites and the strength of purifying selection. This equivalence in neutral diversity patterns suggests that data collected from subdivided populations may have limited power for differentiating among the selective pressures to which closely linked selected sites are subject.     

Rampant host- and defensive phenotype-associated diversification in a goldenrod gall midge

Natural selection can play an important role in the genetic divergence of populations and their subsequent speciation. Such adaptive diversification, or ecological speciation, might underlie the enormous diversity of plant-feeding insects that frequently experience strong selection pressures associated with host plant use as well as from natural enemies. This view is supported by increasing documentation of host-associated (genetic) differentiation in populations of plant-feeding insects using alternate hosts. Here, we examine evolutionary diversification in a single nominal taxon, the gall midge Asteromyia carbonifera (O.S.), with respect to host plant use and gall phenotype. Because galls can be viewed as extended defensive phenotypes of the midges, gall morphology is likely to be a reflection of selective pressures by enemies. Using phylogenetic and comparative analyses of mtDNA and nuclear sequence data, we find evidence that A. carbonifera populations are rapidly diversifying along host plant and gall morphological lines. At a broad scale, geography explains surprisingly little genetic variation, and there is little evidence of strict co-cladogenesis with their Solidago hosts. Gall morphology is relatively labile, distinct gall morphs have evolved repeatedly and colonized multiple hosts, and multiple genetically and morphologically distinct morphs frequently coexist on a single host plant species. These results suggest that Asteromyia carbonifera is in the midst of an adaptive radiation driven by multitrophic selective pressures. Similar complex community pressures are likely to play a role in the diversification of other herbivorous insect groups.     

The role of migration in the evolution of phenotypic switching

Stochastic switching is an example of phenotypic bet hedging, where an individual can switch between different phenotypic states in a fluctuating environment. Although the evolution of stochastic switching has been studied when the environment varies temporally, there has been little theoretical work on the evolution of phenotypic switching under both spatially and temporally fluctuating selection pressures. Here, we explore the interaction of temporal and spatial change in determining the evolutionary dynamics of phenotypic switching. We find that spatial variation in selection is important; when selection pressures are similar across space, migration can decrease the rate of switching, but when selection pressures differ spatially, increasing migration between demes can facilitate the evolution of higher rates of switching. These results may help explain the diverse array of non-genetic contributions to phenotypic variability and phenotypic inheritance observed in both wild and experimental populations.     

Differential migration from high fitness demes in the shining fungus beetle, Phalacrus substriatus

Abstract.— Using data from three years (1994–1996), I tested whether differential migration occurs from demes of high mean fitness in the shining fungus beetle, Phalacrus substriatus . The results show evidence for differential migration, thus providing evidence from a natural population for a critical demographic assumption of many interdemic selection models. To predict the evolutionary response to interdemic selection through differential migration, the genetic basis of the variation among demes in mean fitness must be known because the observed patterns could also be explained by some demes having an intrinsically favorable habitat. Thus, the importance of differential migration through interdemic selection in natural populations cannot be unequivocally answered without experiments specifically addressing the question of what causes differences in mean fitness among demes.     

Interactions Between Entomopathogenic Fungi and Other Natural Enemies: Implications for Biological Control

Pathogens and arthropod natural enemies may contribute to the suppression of insect pest populations either as individual species or as species complexes. However, because natural enemies of insects have evolved and function in a multitrophic context it is important to assess interactions within complexes of natural enemies if they are to be exploited effectively in pest management. Natural enemies can interact either synergistically/additively (e.g. enhanced transmission and dispersal of insect pathogens) or antagonistically (e.g. parasitism/infection, predation and competition). In this paper, studies assessing the potential interactions between insect and fungal natural enemies are reviewed. In general, these studies indicate the positive nature of the interactions between arthropod natural enemies and fungal pathogens with respect to the control of insect populations. More work is required to investigate further the many ways in which the natural enemy community interacts in the agroecosystem     

Spatial heterogeneity in the strength of selection against deleterious alleles and the mutation load

According to current estimates of genomic deleterious mutation rates (which are often of the order 0.1-1) the mutation load (defined as a reduction in the average fitness of a population due to the presence of deleterious alleles) may be important in many populations. In this paper, I use multilocus simulations to explore the effect of spatial heterogeneity in the strength of selection against deleterious alleles on the mutation load (for example, it has been suggested that stressful environments may increase the strength of selection). These simulations show contrasted results: in some situations, spatial heterogeneity may greatly reduce the mutation load, due to the fact that migrants coming from demes under stronger selection carry relatively few deleterious alleles, and benefit from a strong advantage within demes under weaker selection (where individuals carry many more deleterious alleles); in other situations, however, deleterious alleles accumulate within demes under stronger selection, due to migration pressure from demes under weaker selection, leading to fitness erosion within those demes. This second situation is more frequent when the productivity of the different demes is proportional to their mean fitness. The effect of spatial heterogeneity is greatly reduced, however, when the response to environmental differences is inconsistent across loci.     

Putative causes and consequences of MHC variation within and between locally adapted stickleback demes

Genes of the major histocompatibility complex (MHC) have been a source of considerable research interest, owing in large part to the growing body of evidence that they may be subject to both natural and sexual selection. However, much remains to be learned about the dynamics of MHC genes in subdivided populations, particularly those characterized by divergent ecological pressures. In this study, we attempt to disentangle the relative roles of both parasite-mediated selection and MHC-mediated mate choice in an open estuarine system inhabited by two parapatric, adaptively divergent threespine stickleback (Gasterosteus aculeatus) demes. We sequenced the putative peptide-binding region (PBR) of an estimated four Class IIβ loci from 127 individuals, identifying 329 sequence variants (276 translated amino acid sequences). Demes differed significantly both in the frequency of MHC alleles and in the communities of helminth parasites infecting resident sticklebacks. Strong signatures of natural selection were inferred from analyses of codon substitutions, particularly in the derived (freshwater) rather than the ancestral (marine) deme. Relationships between parasite load and MHC diversity were indicative of balancing selection, but only within the freshwater deme. Signals of MHC-mediated mate choice were weak and differed significantly between demes. Moreover, MHC-mediated mate choice was significantly influenced by environmental salinity and appeared of secondary importance to tendencies towards assortative mating. We discuss the implications of these findings in respect to ecological adaptation and the potential demographic consequences of possible outcomes of MHC-mediated mate choice.     

The Distribution of Mutant Alleles in a Subdivided Population

The results are presented from a simulation study of the spatial distribution of mutant alleles in a subdivided population. Statistical measures of the spatial pattern are defined in such a way that the same quantities could be measured in a geographic survey of allele frequencies in natural populations. Two types of quantities are discussed in this paper: (1) the occupancy distribution provides information on the presence or absence of the mutant in different numbers of demes; and (2) the conditional frequency distribution provides information about the extent of local differentiation when the mutant is present in different numbers of demes. Properties of these distributions are found for different types of natural selection acting on the mutant. Some results are presented for the same statistical measures based on samples of individuals from a fraction of the total number of demes. The simulation results for intermediate levels of the migration rates are compared with analytic results obtained on the limits of high and low migration rates. The main conclusion is that these measures of the spatial distribution of mutants in a subdivided population have simple properties that could provide a new perspective on data from natural populations.     

Host-pathogen interactions, insect outbreaks, and natural selection for disease resistance

The theory of insect population dynamics has shown that heterogeneity in natural-enemy attack rates is strongly stabilizing. We tested the usefulness of this theory for outbreaking insects, many of which are attacked by infectious pathogens. We measured heterogeneity among gypsy moth larvae in their risk of infection with a nucleopolyhedrovirus, which is effectively heterogeneity in the pathogen's attack rate. Our data show that heterogeneity in infection risk in this insect is so high that it leads to a stable equilibrium in the models, which is inconsistent with the outbreaks seen in North American gypsy moth populations. Our data further suggest that infection risk declines after epidemics, in turn suggesting that the model assumption of constant infection risk is incorrect. We therefore constructed an alternative model in which natural selection drives fluctuations in infection risk, leading to reductions after epidemics because of selection for resistance and increases after epidemics because of a cost of resistance. This model shows cycles even for high heterogeneity, and experiments confirm that infection risk is indeed heritable. The model is very general, and so we argue that natural selection for disease resistance may play a role in many insect outbreaks.     

Natural selection on fecundity variance in subdivided populations: kin selection meets bet hedging

In a series of seminal articles in 1974, 1975, and 1977, J. H. Gillespie challenged the notion that the "fittest" individuals are those that produce on average the highest number of offspring. He showed that in small populations, the variance in fecundity can determine fitness as much as mean fecundity. One likely reason why Gillespie's concept of within-generation bet hedging has been largely ignored is the general consensus that natural populations are of large size. As a consequence, essentially no work has investigated the role of the fecundity variance on the evolutionary stable state of life-history strategies. While typically large, natural populations also tend to be subdivided in local demes connected by migration. Here, we integrate Gillespie's measure of selection for within-generation bet hedging into the inclusive fitness and game theoretic measure of selection for structured populations. The resulting framework demonstrates that selection against high variance in offspring number is a potent force in large, but structured populations. More generally, the results highlight that variance in offspring number will directly affect various life-history strategies, especially those involving kin interaction. The selective pressures on three key traits are directly investigated here, namely within-generation bet hedging, helping behaviors, and the evolutionary stable dispersal rate. The evolutionary dynamics of all three traits are markedly affected by variance in offspring number, although to a different extent and under different demographic conditions.     

Migration promotes mutator alleles in subdivided populations

Mutator alleles that elevate the genomic mutation rate may invade nonrecombining populations by hitchhiking with beneficial mutations. Mutators have been repeatedly observed to take over adapting laboratory populations and have been found at high frequencies in both microbial pathogen and cancer populations in nature. Recently, we have shown that mutators are only favored by selection in sufficiently large populations and transition to being disfavored as population size decreases. This population size‐dependent sign inversion in selective effect suggests that population structure may also be an important determinant of mutation rate evolution. Although large populations may favor mutators, subdividing such populations into sufficiently small subpopulations (demes) might effectively inhibit them. On the other hand, migration between small demes that otherwise inhibit hitchhiking may promote mutator fixation in the whole metapopulation. Here, we use stochastic, agent‐based simulations and evolution experiments with the yeast Saccharomyces cerevisiae to show that mutators can, indeed, be favored by selection in subdivided metapopulations composed of small demes connected by sufficient migration. In fact, we show that population structure plays a previously unsuspected role in promoting mutator success in subdivided metapopulations when migration is rare.     

Diffuse selection on resistance to deer herbivory in the ivyleaf morning glory, Ipomoea hederacea

Recent work defines coevolution between plants and herbivores as pairwise when the pattern of selection on resistance traits and the response to selection are both independent of the presence or absence of other herbivores. In addition, for a pairwise response to selection, resistance to a focal herbivore must have the same genetic basis in the presence and absence of other herbivores. None of these conditions were satisfied for the ivyleaf morning glory, Ipomoea hederacea, and its insect, fungal, and mammalian natural enemies with a quantitative genetics field experiment. A significant negative genetic correlation exists between resistance to deer and generalist insect herbivory that would preclude an independent response to selection. In addition, resistance loci under selection differ depending on the composition of the natural enemy community as indicated by genetic correlations between deer resistances in the presence and absence of other natural enemies that differ substantially from 1. Finally, selection on deer resistance depends on the presence or absence of insects; in the presence of insects, greater deer resistance is favored, but in the absence of insects, deer resistance is effectively neutral. These results indicate that the composition of the natural enemy community can alter both the pattern of selection and the likely response to selection of resistance traits.     

昆虫对环境低氧的适应:生理和分子机制研究进展

氧是机体进行新陈代谢和维持生存的必要因素。低氧环境在自然界普遍存在,也是许多重大疾病(如癌症)发生过程中基本的病理生理特征。生物包括昆虫在其生存和发育过程中经常面对低氧的挑战,它们发展出了各自的适应策略以求得生存和繁荣壮大。昆虫对于低氧环境适应包括在气管系统通气量、气体交换模式、体型大小和发育时间等生理机制上的改变。为揭示昆虫低氧适应机制,研究人员针对不同昆虫采用了来自人工选择或者自然选择的品系(种群),使用了基因芯片表达和转录组测序、基因组重测序技术和基因操作等技术。基于这些方法研究发现,在分子机制方面,昆虫可以通过抑制能量代谢、提高氧气利用率来适应低氧环境;还可以通过胰岛素通路、低氧诱导因子(HIF)信号通路等来调节自身代谢活动从而适应环境低氧;除此之外,昆虫的气管系统可以在基因调控下通过代偿性生理和形态变化来适应低氧环境。昆虫低氧适应机制的研究为探求昆虫数亿年进化过程中体形改变、物种形成、种群动态等提供提供新的视野,也增进对动物应对低氧或缺氧机理的深入理解,特别是为研究人类重大疾病的发生提供重要启示。     

Scale insect allozyme differentiation within and between host trees

Summary Allelic frequencies and genotypic distributions in three polymorphic enzyme systems demonstrated genetic differentiation over extraordinarily short distances in a population of black pineleaf scale insects infesting ponderosa pine trees. A hierarchical analysis of the population genetic structure showed significant differences between demes on different twigs within individual host trees, between demes on neighboring trees, and between demes in pine plots on adjacent city blocks. Allelic frequencies at a malic enzyme locus were associated with deme-to-deme variation in ecological correlates of insect fitness, suggesting adaptive hypotheses about the causes of population subdivision.     

Host range in phytophagous insects: the potential role of generalist predators

Summary The potential role of generalist natural enemies is presented as one of the important ecological pressures that select for narrow host range in phytophagous insects, and dominant relative to physiological bases for specialization. Experiments are described in three completely different systems indicating that generalist herbivores are more vulnerable to predation than specialist herbivores. The three predators were (a) the vespid waspMischocyttarus flavitarsus, (b) the Argentine antIridomyrmex humilis and (c) the coccinellid beetleHippodamia convergens. It is concluded the predators may provide strong selection pressure for maintenance and perhaps evolution of narrow host range in insect herbivores.     

Population genetics in the American Tropics XVI. Data on partial dominance of recessives in Drosophila willistoni

Summary Through a series of genetic load studies made on 1) samples of Drosophila willistoni from two sites in Mesitas, Colombia, it was found that the relative contributions to the total, subvital and lethal loads reflect lethal equivalences (B/A) ratios which support more the balancing theory of population structure than the neutralist theory. Moreover, measurements of population size have revealed the existance of very small demes in local populations. Under such conditions we have calculated extremely small lethal equivalence ratios in demes where probably a great deal of consanguinity takes place. We are aware that under these conditions B/A ratios cannot be very good monitors of random load measurements and, therefore, suggest a change in the mathematical formulation that take into consideration the existance of small populations.Furthermore, it appears plausible that the degree of penetrance in the heterozygous condition changes as the population structure changes. We speculate that natural populations may have unknown selective mechanisms capable of guiding unknown dominance modifiers according to the intensity of selection.     

Genealogy of Neutral Genes and Spreading of Selected Mutations in a Geographically Structured Population

In a geographically structured population, the interplay among gene migration, genetic drift and natural selection raises intriguing evolutionary problems, but the rigorous mathematical treatment is often very difficult. Therefore several approximate formulas were developed concerning the coalescence process of neutral genes and the fixation process of selected mutations in an island model, and their accuracy was examined by computer simulation. When migration is limited, the coalescence (or divergence) time for sampled neutral genes can be described by the convolution of exponential functions, as in a panmictic population, but it is determined mainly by migration rate and the number of demes from which the sample is taken. This time can be much longer than that in a panmictic population with the same number of breeding individuals. For a selected mutation, the spreading over the entire population was formulated as a birth and death process, in which the fixation probability within a deme plays a key role. With limited amounts of migration, even advantageous mutations take a large number of generations to spread. Furthermore, it is likely that these mutations which are temporarily fixed in some demes may be swamped out again by non-mutant immigrants from other demes unless selection is strong enough. These results are potentially useful for testing quantitatively various hypotheses that have been proposed for the origin of modern human populations.     

Local maladaptation in the soft scale insect Saissetia coffeae (Hemiptera: Coccidae)

Local adaptation has often been documented in herbivorous insects. The potential for local maladaptation in phytophagous insects, however, has not been widely considered. I performed a two-generation reciprocal cross-transplant experiment with the generalist soft scale insect Saissetia coffeae (Hemiptera: Coccidae) on two common species of host plants in rain forest habitat in Costa Rica. In this system, S. coffeae showed significant local maladaptation at the level of the host species. Lineages originally collected from Witheringia enjoyed a strong advantage over those collected from Lomariopsis when both sets of lineages were placed on Lomariopsis; however, when both sets of lineages were raised on Witheringia, their fitnesses were statistically indistinguishable. While some aspects of the biology of S. coffeae may impair its ability to adapt to local selection pressures, scale insects are often locally adapted on fine spatial scales, and local maladaptation is therefore especially surprising. Other documented cases of local maladaptation in parasites appear to be due to evolution on the part of the host. The possibility that hosts or natural enemies may place local genotypes at a disadvantage, producing a pattern of local maladaptation, is one that deserves more consideration in the context of plant-insect interactions.     

Reducing the impact of insect herbivory in eucalypt plantations through management of extrinsic influences on tree vigour

In this paper it is argued that concepts developed in ecologically derived insect–plant interaction models can contribute directly to the management of insect herbivory in eucalypt plantations. Common to most species of commercially planted eucalypt is their genetic potential for early rapid growth. Several plant defence theories predict that intrinsically fast growing plants are able to tolerate relatively high levels of herbivory. The risk of this strategy failing increases when plants are exposed to external stressful factors that reduce canopy growth and vigour. Results from a young Eucalyptus camaldulensis plantation stressed by moisture deficit and two young Eucalyptus dunnii plantations, stressed by flooding and weed competition, respectively, are summarized. In all three cases, the stress‐inducing agents reduced canopy growth rates and architecture so that the proportion of leaf tissue damaged by insects increased and the tree’s ability to tolerate this damage decreased. Therefore, alleviating tree stress through improved silvicultural practices or improved site selection techniques may indirectly reduce the impact of insect herbivory. In resource‐limiting environments, an alternative approach may be to plant eucalypt species that although slower growing, are predicted to have better defended foliage. Manipulation of these natural antiherbivore plant strategies are not exclusive of other management approaches, such as the need for routine surveillance of key pest insects or the genetic selection of natural insect resistance and selective chemical control techniques, but should be viewed as an overarching concept for plantation health.