Do mites evolving in alternating host plants adapt to host switch?

A fluctuating environment may be perceived as a composition of different environments, or as an environment per se, in which it is the fluctuation itself that poses a selection pressure. If so, then organisms may adapt to this alternation. We tested this using experimental populations of spider mites that have been evolving for 45 generations in a homogeneous environment (pepper or tomato plants), or in a heterogeneous environment composed of an alternation of these two plants approximately at each generation. The performance (daily oviposition rate and juvenile survival) of individuals from these populations was tested in each of the homogeneous environments, and in two alternating environments, one every 3 days and the other between generations. To discriminate between potential genetic interactions between alleles conferring adaptation to each host plant and environmental effects of evolving in a fluctuating environment, we compared the performance of all lines with that of a cross between tomato and pepper lines. As a control, two lines within each selection regime were also crossed. We found that crosses between alternating lines and between pepper and tomato lines performed worse than crosses between lines evolving in homogeneous environments when tested in that environment. In contrast, alternating lines performed either better or similarly to lines evolving in homogeneous environments when tested in a fluctuating environment. Our results suggest that fluctuating environments are more than the juxtaposition of two environments. Hence, tests for adaptation of organisms evolving in such environments should be carried out in fluctuating conditions.     

Heritable variation in host tolerance and resistance inferred from a wild host–parasite system

Hosts have evolved two distinct defence strategies against parasites: resistance (which prevents infection or limit parasite growth) and tolerance (which alleviates the fitness consequences of infection). However, heritable variation in resistance and tolerance and the genetic correlation between these two traits have rarely been characterized in wild host populations. Here, we estimate these parameters for both traits in Leuciscus burdigalensis, a freshwater fish parasitized by Tracheliastes polycolpus. We used a genetic database to construct a full-sib pedigree in a wild L. burdigalensis population. We then used univariate animal models to estimate inclusive heritability (i.e. all forms of genetic and non-genetic inheritance) in resistance and tolerance. Finally, we assessed the genetic correlation between these two traits using a bivariate animal model. We found significant heritability for resistance (H = 17.6%; 95% CI: 7.2–32.2%) and tolerance (H = 18.8%; 95% CI: 4.4–36.1%), whereas we found no evidence for the existence of a genetic correlation between these traits. Furthermore, we confirm that resistance and tolerance are strongly affected by environmental effects. Our results demonstrate that (i) heritable variation exists for parasite resistance and tolerance in wild host populations, and (ii) these traits can evolve independently in populations.     

Coevolution of dispersal in a parasitoid–host system

Interspecific interactions and the evolution of dispersal are both of interest when considering the potential impact of habitat fragmentation on community ecology, but the interaction between these processes is not well studied. We address this by considering the coevolution of dispersal strategies in a host–parasitoid system. An individual-based host–parasitoid metapopulation model was constructed for a patchy environment, allowing for evolution in dispersal rates of both species. Highly rarefied environments with few suitable patches selected against dispersal in both species, as did relatively static environments. Provided that parasitoids persist, all the variables studied led to stable equilibria in dispersal rates for both species. There was a tendency toward higher dispersal rates in parasitoids because of the asymmetric relationships of the two species to the patches: vacant patches are most valuable for hosts, but unsuitable for parasitoids, which require an established host population to reproduce. High host dispersal rate was favoured by high host population growth rate, and in the parasitoid by high growth rates in both species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Environment–host–parasite interactions in mass-reared insects

The mass production of insects is rapidly expanding globally, supporting multiple industrial needs. However, parasite infections in insect mass-production systems can lower productivity and can lead to devastating losses. High rearing densities and artificial environmental conditions in mass-rearing facilities affect the insect hosts as well as their parasites. Environmental conditions such as temperature, gases, light, vibration, and ionizing radiation can affect productivity in insect mass-production facilities by altering insect development and susceptibility to parasites. This review explores the recent literature on environment–host–parasite interactions with a specific focus on mass-reared insect species. Understanding these complex interactions offers opportunities to optimise environmental conditions for the prevention of infectious diseases in mass-reared insects.     

Climatic effects and phenological mismatch in cuckoo–host interactions: a role for host phenotypic plasticity in laying date?

Climatic effects on breeding phenology vary across organisms and therefore might promote a phenological mismatch in ecologically interacting species, including those engaged in coevolutionary interactions such as brood parasites and their hosts. Recent studies suggest that climatic induced changes in migration phenology may have mismatched cuckoos and their hosts in Europe. However, it is currently unknown whether cuckoo–host phenological mismatch results from different degrees of phenotypic plasticity or to different speeds of microevolutionary processes affecting hosts and parasites. Here we performed 1) cross‐sectional correlations between climate conditions and population level of phenological mismatch between the migratory brood parasite great spotted cuckoo Clamator glandarius and its main resident host in Europe, the magpie Pica pica; and 2) a longitudinal analysis to study within‐individual variation in breeding phenology for individual hosts experiencing different climate conditions over a period of nine years (2005–2013). Cross‐sectional analyses revealed independent and contrary effects of winter and spring temperature on magpie phenology: magpie hosts tend to breed earlier those years with lower February temperatures, however, high temperature in the first half of April spur individuals to lay eggs. Breeding phenology of cuckoos was tuned to that of their magpie host in time and duration. However, annual phenological mismatch between cuckoos and magpie hosts increased with NAO index and January temperature. Longitudinal analyses revealed high individual consistency in magpie host phenology, but a low influence of climate, suggesting that the climatic‐driven phenological mismatch between cuckoos and magpies at the population‐level cannot be explained by a host plastic response to climatic conditions.     

Addicted? Reduced host resistance in populations with defensive symbionts

Heritable symbionts that protect their hosts from pathogens have been described in a wide range of insect species. By reducing the incidence or severity of infection, these symbionts have the potential to reduce the strength of selection on genes in the insect genome that increase resistance. Therefore, the presence of such symbionts may slow down the evolution of resistance. Here we investigated this idea by exposing Drosophila melanogaster populations to infection with the pathogenic Drosophila C virus (DCV) in the presence or absence of Wolbachia, a heritable symbiont of arthropods that confers protection against viruses. After nine generations of selection, we found that resistance to DCV had increased in all populations. However, in the presence of Wolbachia the resistant allele of pastrel—a gene that has a major effect on resistance to DCV—was at a lower frequency than in the symbiont-free populations. This finding suggests that defensive symbionts have the potential to hamper the evolution of insect resistance genes, potentially leading to a state of evolutionary addiction where the genetically susceptible insect host mostly relies on its symbiont to fight pathogens.     

Isotopic insight into host–endosymbiont relationships in Liolaemid lizards

Nitrogen isotopes have been widely used to investigate trophic levels in ecological systems. Isotopic enrichment of 2–5‰ occurs with trophic level increases in food webs. Host–parasite relationships deviate from traditional food webs in that parasites are minimally enriched relative to their hosts. Although this host–parasite enrichment pattern has been shown in multiple systems, few studies have used isotopic relationships to examine other potential symbioses. We examined the relationship between two gut-nematodes and their lizard hosts. One species, Physaloptera retusa, is a documented parasite in the stomach, whereas the relationship of the other species, Parapharyngodon riojensis (pinworms), to the host is putatively commensalistic or mutualistic. Based on the established trophic enrichments, we predicted that, relative to host tissue, parasitic nematodes would be minimally enriched (0–1‰), whereas pinworms, either as commensals or mutualists, would be significantly enriched by 2–5‰. We measured the ¹⁵N values of food, digesta, gut tissue, and nematodes of eight lizard species in the family Liolaemidae. Parasitic worms were enriched 1±0.2‰ relative to host tissue, while the average enrichment value for pinworms relative to gut tissue was 6.7±0.2‰. The results support previous findings that isotopic fractionation in a host–parasite system is lower than traditional food webs. Additionally, the larger enrichment of pinworms relative to known parasites suggests that they are not parasitic and may be several trophic levels beyond the host.     

Is the host or the parasitoid in control?: effects of host age and temperature on pseudoparasitization by Microplitis rufiventris in Spodoptera littoralis

We compared the production of pseudoparasitization by Microplitis rufiventris females in most (third) and less (fourth) preferred instars of Spodoptera littoralis larvae at 20+/-1 and 27+/-1 degrees C. The parasitized hosts were classified into hosts producing parasitoids (type A hosts) and hosts producing no parasitoids, i.e., pseudoparasitized hosts (type B hosts). The latter were further classified into: (a) pseudoparasitized hosts with "well" arrested development (type B1 hosts); (b) pseudoparasitized hosts with partially arrested development (type B2 hosts); and (c) pseudoparasitized hosts that successfully pupated to apparently normal host pupae (type B3 hosts). The present series of experiments showed that parasitization by M. rufiventris was clearly affected by host instar, age within an instar and rearing temperature. Production of type B hosts was less when third instar S. littoralis larvae were exposed to the wasp females than when the host larvae were in fourth instar. The production of type A hosts was much greater when early or mid ages of an instar was stung by the wasp females comparing with stung late age of the same instar. Production of type B hosts may be due to one or overall of the following: (a) dosage dilution of M. rufiventris female's factors in the different age classes of the instar; (b) endocrine system (physiological state) at parasitization time, i.e., early vs late age of the instar; (c) growth rate of host larvae. The lowest production of type B hosts was at highest growth rate; and (d) temperature, larger proportions of type B hosts were produced at 27+/-1 than at 20+/-1degrees C. The three types host development (B1, B2 and B3) are possibly representing three levels of host resistance (host control) resulting in partial or complete failure of parasitoid control. Type A hosts represent complete success of parasitoid control. The results suggest that the impact of parasitoid factor(s) on developmental arrest is affected by host age at the time of parasitism and/or by temperature.     

The scaling of parasite biomass with host biomass in lake ecosystems: are parasites limited by host resources?

The standing crop biomass of different populations or trophic levels reflects patterns of energy flow through an ecosystem. The contribution of parasites to total biomass is often considered negligible; recent evidence suggests otherwise, although it comes from a narrow range of natural systems. Quantifying how local parasite biomass, whether that of a single species or an assemblage of species sharing the same host, varies across localities with host population biomass, is critical to determine what constrains parasite populations. We use an extensive dataset on all free‐living and parasitic metazoan species from multiple sites in New Zealand lakes to measure parasite biomass and test how it covaries with host biomass. In all lakes, trematodes had the highest combined biomass among parasite taxa, ranging from about 0.01 to 0.25 g m^?2, surpassing the biomass of minor free‐living taxa. Unlike findings from other studies, the life stage contributing the most to total trematode biomass was the metacercarial stage in the second intermediate host, and not sporocysts or rediae within snail first intermediate hosts, possibly due to low prevalence and small snail sizes. For populations of single parasite species, we found no relationship between host and parasite biomass for either juvenile or adult nematodes. In contrast, all life stages of trematodes had local biomasses that correlated positively with those of their hosts. For assemblages of parasite species sharing the same host, we found strong relationships between local host population biomass and the total biomass of parasites supported. In these host–parasite biomass relationships, the scaling factor (slope in log‐log space) suggests that parasites may not be making full use of available host resources. Host populations appear capable of supporting a little more parasite biomass, and may be open to expansion of existing parasites or invasion by new ones.     

A host–microparasite model with a resistant host

The theory of heterozygote advantage is often used to explain the genetic variation found in natural populations. If a large population randomly mates and the various genotypes have the same growth and death rates, the evolution of the genotypes follows Hardy–Weinberg proportions and polymorphism results. When other environmental stresses, like predators, prey and diseases, are present, polymorphism may or may not occur depending on how the various genotypes are affected by the stress. In this paper, we use a basic host–microparasite model to demonstrate that polymorphism can occur even if one genotype suffers a higher death rate than the others in the absence of the parasite if the heterozygote has resistance or immunity to the parasite.     

Population structure of Spodoptera frugiperda maize and rice host forms in South America: are they host strains?

Determining which factors contribute to the formation and maintenance of genetic divergence to evaluate their relative importance as a cause of biological differentiation is among the major challenges in evolutionary biology. In Spodoptera frugiperda (Smith) (Lepidoptera: Noctuidae) two host strains have been recognized in the 1980s: the corn‐strain prefers maize, sorghum, and cotton, whereas the rice‐strain prefers rice and wild grasses. However, it is not clear to what extent these so‐called ‘strains’, which have also been called ‘host races’ or even ‘sibling species’, are really associated with host plants. Due to the indeterminate evolutionary status, we will use the term ‘host forms’ (sensu Funk). Here, we characterized populations collected from maize, rice, and wild grasses from three countries in South America. Using two mitochondrial cytochrome oxidase I (mtCOI) markers and 10 polymorphisms in the triose phosphate isomerase (Tpi) gene, we found various patterns of host association. Two hundred twenty‐seven nuclear amplified fragment length polymorphisms (AFLPs) markers revealed significant genetic differentiation among populations, which was generally correlated to the host from which the larvae were collected. Using a multivariate discriminant analysis and a Bayesian clustering approach, we found that individuals could be grouped into 2–5 genetically distinct clusters, depending on the method. Together, our results indicate that although host‐associated differentiation is present in this species, it does not account for all observable genetic variation and other factors must be maintaining genetic differentiation between these forms. Therefore, the term ‘host strains’ should be abandoned and ‘host forms’ should be used instead for S. frugiperda.     

Can imperfect host discrimination explain partial patch exploitation in parasitoids?

1. Host discrimination by Aphidius rhopalosiphi (De Stefani Perez) (Hymenoptera: Braconidae) was first studied on the grain aphid Sitobion avenae (Fabricius) (Homoptera: Aphididae). Females tended to avoid oviposition in hosts parasitised 3 h earlier. No evidence of host discrimination ability on freshly parasitised hosts was suggested, however, and ovipositional experience had no effect on host discrimination. 2. The effects of host discrimination ability on the exploitation strategy of patches containing different proportions of unparasitised hosts and hosts parasitised for 3 h were studied. Females spent less time on patches with a higher proportion of parasitised hosts, reflecting the females' ability to perceive the potential profitability of the patch. This ability may be based on the nature of the hosts encountered (unparasitised or parasitised). 3. Incomplete exploitation of unparasitised hosts was also observed. It seems that this partial exploitation is related to the inability of A. rhopalosiphi to recognise freshly parasitised hosts. As a female may experience a risk of self‐superparasitism during patch depletion, this could promote early departure from incompletely exploited patches. 4. The effect of previous experience on the patch exploitation strategy was also assessed. Females were tested twice on two patches of the same quality. Results suggested that the experience acquired during a previous visit led the females to leave the patch sooner and to lay fewer eggs in parasitised hosts. 5. Patch exploitation strategy may therefore be the result of different factors such as host discrimination and experience. The evolutionary consequences of the results are discussed.     

Stochastic environmental fluctuations drive epidemiology in experimental host–parasite metapopulations

Environmental fluctuations are important for parasite spread and persistence. However, the effects of the spatial and temporal structure of environmental fluctuations on host–parasite dynamics are not well understood. Temporal fluctuations can be random but positively autocorrelated, such that the environment is similar to the recent past (red noise), or random and uncorrelated with the past (white noise). We imposed red or white temporal temperature fluctuations on experimental metapopulations of Paramecium caudatum, experiencing an epidemic of the bacterial parasite Holospora undulata. Metapopulations (two subpopulations linked by migration) experienced fluctuations between stressful (5°C) and permissive (23°C) conditions following red or white temporal sequences. Spatial variation in temperature fluctuations was implemented by exposing subpopulations to the same (synchronous temperatures) or different (asynchronous temperatures) temporal sequences. Red noise, compared with white noise, enhanced parasite persistence. Despite this, red noise coupled with asynchronous temperatures allowed infected host populations to maintain sizes equivalent to uninfected populations. It is likely that this occurs because subpopulations in permissive conditions rescue declining subpopulations in stressful conditions. We show how patterns of temporal and spatial environmental fluctuations can impact parasite spread and host population abundance. We conclude that accurate prediction of parasite epidemics may require realistic models of environmental noise.     

Does genetic diversity limit disease spread in natural host populations?

It is a commonly held view that genetically homogenous host populations are more vulnerable to infection than genetically diverse populations. The underlying idea, known as the 'monoculture effect,' is well documented in agricultural studies. Low genetic diversity in the wild can result from bottlenecks (that is, founder effects), biparental inbreeding or self-fertilization, any of which might increase the risk of epidemics. Host genetic diversity could buffer populations against epidemics in nature, but it is not clear how much diversity is required to prevent disease spread. Recent theoretical and empirical studies, particularly in Daphnia populations, have helped to establish that genetic diversity can reduce parasite transmission. Here, we review the present theoretical work and empirical evidence, and we suggest a new focus on finding 'diversity thresholds.'     

Symbiont genes in host genomes: fragments with a future?

While lateral transfer is the rule in the evolutionary history of bacterial and archaeal genes, events of transfer from prokaryotes to eukaryotes are rare. Germline-transmitted animal symbionts, such as Wolbachia pipientis, are well placed to participate in such transfers. In a recent issue of Science, Dunning Hotopp et al. identified instances of transfer of Wolbachia DNA to host genomes. It is unknown whether these transfers represent innovation in animal evolution.     

Delayed feedback and multiple attractors in a host–parasitoid system

 Continuous-time, age structured, host–parasitoid models exhibit three types of cyclic dynamics: Lotka–Volterra-like consumer-resource cycles, discrete generation cycles, and “delayed feedback cycles” that occur if the gain to the parasitoid population (defined by the number of new female parasitoid offspring produced per host attacked) increases with the age of the host attacked. The delayed feedback comes about in the following way: an increase in the instantaneous density of searching female parasitoids increases the mortality rate on younger hosts, which reduces the density of future older and more productive hosts, and hence reduces the future per head recruitment rate of searching female parasitoids. Delayed feedback cycles have previously been found in studies that assume a step-function for the gain function. Here, we formulate a general host–parasitoid model with an arbitrary gain function, and show that stable, delayed feedback cycles are a general phenomenon, occurring with a wide range of gain functions, and strongest when the gain is an accelerating function of host age. We show by examples that locally stable, delayed feedback cycles commonly occur with parameter values that also yield a single, locally stable equilibrium, and hence their occurrence depends on initial conditions. A simplified model reveals that the mechanism responsible for the delayed feedback cycles in our host–parasitoid models is similar to that producing cycles and initial-condition-dependent dynamics in a single species model with age-dependent cannibalism. Received: 24 October 1997 / Revised version: 13 June 1998