Effect of temperature and host-parasite ratio on sex differentiation of Romanomermis iyengari (Welch), a mermithid parasite of mosquitoes

The effect of temperature and host-parasite ratio on the percentage infection and sex differentiation of R. iyengari was studied. Significant differences were observed in the percentage infection due to different host-parasite ratios and temperatures. At 25 degrees and 30 degrees C, the host parasite ratio of 1:3 resulted in 86-92% infection of Culex quinquefasciatus larvae. At 20 degrees and 35 degrees C, a higher host-parasite ratio was required to get this level of infection. More number of post-parasites per mosquito larva emerged at 20 degrees (1.5-5.8) and 25 degrees C (1.9-6.3) than at 30 degrees (1.5-3.9) and 35 degrees C (1.6-3.6). More than 50% of the post-parasites were females at 20 degrees and 25 degrees, 30 degrees and 35 degrees C at 1:1-1:10, 1:1-1:4 and 1:1-1:3 host-parasite ratios, respectively.     

Divergent Macroparasite Infections in Parapatric Swiss Lake-Stream Pairs of Threespine Stickleback (Gasterosteus aculeatus)

Spatial heterogeneity in diversity and intensity of parasitism is a typical feature of most host-parasite interactions, but understanding of the evolutionary implications of such variation is limited. One possible outcome of infection heterogeneities is parasite-mediated divergent selection between host populations, ecotypes or species which may facilitate the process of ecological speciation. However, very few studies have described infections in population-pairs along the speciation continuum from low to moderate or high degree of genetic differentiation that would address the possibility of parasite-mediated divergent selection in the early stages of the speciation process. Here we provide an example of divergent parasitism in freshwater fish ecotypes by examining macroparasite infections in threespine stickleback (Gasterosteus aculeatus) of four Swiss lake systems each harbouring parapatric lake-stream ecotype pairs. We demonstrate significant differences in infections within and between the pairs that are driven particularly by the parasite taxa transmitted to fish from benthic invertebrates. The magnitude of the differences tended to correlate positively with the extent of neutral genetic differentiation between the parapatric lake and stream populations of stickleback, whereas no such correlation was found among allopatric populations from similar or contrasting habitats. This suggests that genetic differentiation is unrelated to the magnitude of parasite infection contrasts when gene flow is constrained by geographical barriers while in the absence of physical barriers, genetic differentiation and the magnitude of differences in infections tend to be positively correlated.     

Temperature-dependent transmission and latency of Holospora undulata, a micronucleus-specific parasite of the ciliate Paramecium caudatum

Transmission of parasites to new hosts crucially depends on the timing of production of transmission stages and their capacity to start an infection. These parameters may be influenced by genetic factors, but also by the environment. We tested the effects of temperature and host genotype on infection probability and latency in experimental populations of the ciliate Paramecium caudatum, after exposure to infectious forms of its bacterial parasite Holospora undulata. Temperature had a significant effect on the expression of genetic variation for transmission and maintenance of infection. Overall, low temperature (10 degrees C) increased levels of (multiple) infection, but arrested parasite development; higher temperatures (23 and 30 degrees C) accelerated the onset of production of infectious forms, but limited transmission success. Viability of infectious forms declined rapidly at 23 and 30 degrees C, thereby narrowing the time window for transmission. Thus, environmental conditions can generate trade-offs between transmission relevant parameters and alter levels of multiple infection or parasite-mediated selection, which may affect evolutionary trajectories of parasite life history or virulence.     

Food-environment mediates the outcome of specific interactions between a bumblebee and its trypanosome parasite

Specific host-parasite interactions, where the outcome of exposure to a parasite depends upon the genotypic identity of both parties, have implications for understanding host-parasite coevolution and patterns of genetic diversity. Thus, grasping the extent to which these interactions are mediated by environmental changes in a spatially and temporally heterogeneous world is vital. In this study, it is shown that the environment can influence specific host-parasite interactions in the well-studied system of the bumblebee Bombus terrestris and its trypanosome parasite Crithidia bombi. Naturally relevant variation in the quality of the food environment formed a three-way interaction with both host and parasite identity in determining the outcome of infection, with regard to the resistance of the host and the transmission of the parasite. The demonstration of such a host-genotype by parasite-genotype by environment interaction (G(H) x G(P) x E) shows the importance of considering environmental variation when investigating host-parasite interactions. Moreover, such interactions may to some extent explain levels of genetic diversity in natural host-parasite systems owing to the fact that they will create selection mosaics when environments are heterogeneous.     

Parasite-mediated selection and the role of sex and diapause in Daphnia

To gain insight into parasite-mediated natural selection, we studied a natural population of the crustacean Daphnia magna during a severe epidemic of the bacterial parasite Pasteuria ramosa. We also investigated the relationship between susceptibility and the production of resting eggs, which are only produced during the sexual phase of reproduction. Live host samples were taken before and after this epidemic and resistance to P. ramosa was examined in the laboratory. Host clones collected after the epidemic were more resistant to P. ramosa than were those collected pre-epidemic, which is consistent with parasite-mediated selection. In our study population, asexually reproducing females were observed across the entire study period, but females carrying resting eggs were observed only prior to the epidemic. For hosts isolated in this pre-epidemic period, we found evidence that those carrying resting eggs (at the time of collection) were more susceptible than those that were reproducing asexually. This was especially apparent for measures of parasite growth, although not all measures of infection success conclusively supported this pattern. Nevertheless, the data suggest that some genotypes invest heavily in diapause at the expense of immunocompetence. Sex could therefore inhibit the evolution of resistance because each spring new genotypes will hatch from resting eggs that are relatively susceptible as they were not exposed to the previous years bout of parasite-mediated selection.     

Sexual selection and the geography of Plasmodium infection in Savannah sparrows (Passerculus sandwichensis)

According to Hamilton and Zuk's hypothesis of parasite-mediated sexual selection, host-parasite coevolution maintains variation in male genetic quality and allows for strong intersexual selection in species with high rates of infection. In birds, most interspecific tests of this hypothesis relate the prevalence of blood parasites to some measure of the intensity of sexual selection. Such tests often rely on limited sampling of single populations to estimate species-wide infection rates, and many tests are thus vulnerable to intraspecific (geographic) variation in the evolutionary ecology of disease. Here, we used molecular techniques to examine variation in the prevalence of Plasmodium spp. across 14 populations of Savannah sparrows, Passerculus sandwichensis, in eastern North America. Plasmodium could not be detected in any of 68 island birds, but 34 of 119 (29%) mainland males, and 7 of 43 (16%) mainland females were infected. Among mainland birds, infection was common in southern populations but rare in New Brunswick, Canada. Overall, the prevalence of Plasmodium ranged from 0 to 60% across populations, although only 17.8% of birds were infected in the pooled (species-wide) sample. The extent of this geographic variation suggests that limited sampling of single populations is unlikely to yield accurate estimates of species-wide infection rates. However, among mainland Savannah sparrows, the prevalence of malaria correlated strongly with average male size and the degree of sexual size dimorphism. We speculate that either sexual selection leads to male-biased infection or, conversely, that high rates of infection promote the evolution of strong intersexual selection.     

Parasite-mediated disruptive selection in a natural <Emphasis Type="Italic">Daphnia</Emphasis> population

Background  

A mismatch has emerged between models and data of host-parasite evolution. Theory readily predicts that parasites can promote host diversity through mechanisms such as disruptive selection. Yet, despite these predictions, empirical evidence for parasite-mediated increases in host diversity remains surprisingly scant.     

Parasite diversity,patterns of MHC II variation and olfactory based mate choice in diverging three-spined stickleback ecotypes

Ecological speciation has been the subject of intense research in evolutionary biology but the genetic basis of the actual mechanism driving reproductive isolation has rarely been identified. The extreme polymorphism of the major histocompatibility complex (MHC), probably maintained by parasite-mediated selection, has been proposed as a potential driver of population divergence. We performed an integrative field and experimental study using three-spined stickleback river and lake ecotypes. We characterized their parasite load and variation at MHC class II loci. Fish from lakes and rivers harbor contrasting parasite communities and populations possess different MHC allele pools that could be the result of a combined action of genetic drift and parasite-mediated selection. We show that individual MHC class II diversity varies among populations and is lower in river ecotypes. Our results suggest the action of homogenizing selection within habitat type and diverging selection between habitat types. Finally, reproductive isolation was suggested by experimental evidence: in a flow channel design females preferred assortatively the odor of their sympatric male. This demonstrates the role of olfactory cues in maintaining reproductive isolation between diverging fish ecotypes.     

Geographic and within-population structure in variable resistance to parasite species and strains in a vertebrate host

Host resistance to parasites and parasite infectivity may be subject to significant genetically determined variation within species. However, relatively little is known of how this variability is structured in natural vertebrate populations and their macroparasites. A laboratory experiment on host susceptibility-parasite infectivity variation in a wildlife host-parasite system (subspecies of the anuran X. laevis and their polystome flatworms), including 33 pairwise allopatric and sympatric host-parasite combinations (three parasite geographical isolates x 11 host full-sibling families, n=600), revealed a complex pattern of infection success. Results amongst host sibships from different localities suggested that infection success was subject to a highly significant locality x parasite isolate interaction. Within localities, a highly significant sibship x isolate interaction also occurred in one of two groups of sibships examined. The existence of such interactions suggests a potential for frequency-dependent, Red Queen-like selection. Interaction between locality and isolate was partly due to higher infection levels in sympatric combinations, consistent with a general pattern of host-specific adaptation. However, some allopatric combinations produced unpredictably high infection levels, resulting in very asymmetrical cross-infectivity patterns (where the reciprocal cross-infections produced negligible infection). This phylogeographically structured host-parasite system may, therefore, sometimes generate local parasite strains with high infectivity to allopatric hosts. Secondary contact between populations could thus result in significant, and unequal, transfer of parasites.     

Parasitism, host immune function, and sexual selection

Parasite-mediated sexual selection may arise as a consequence of 1) females avoiding mates with directly transmitted parasites, 2) females choosing less-parasitized males that provide parental care of superior quality, or 3) females choosing males with few parasites in order to obtain genes for parasite resistance in their offspring. Studies of specific host-parasite systems and comparative analyses have revealed both supportive and conflicting evidence for these hypotheses. A meta-analysis of the available evidence revealed a negative relationship between parasite load and the expression of male secondary sexual characters. Experimental studies yielded more strongly negative relationships than observations did, and the relationships were more strongly negative for ectoparasites than for endoparasites. There was no significant difference in the magnitude of the negative effect for species with and without male parental care, or between behavioral and morphological secondary sexual characters. There was a significant difference between studies based on host immune function and those based on parasite loads, with stronger effects for measures of immune function, suggesting that the many negative results from previous analyses of parasite-mediated sexual selection may be explained because relatively benign parasites were studied. The multivariate analyses demonstrating strong effect sizes of immune function in relation to the expression of secondary sexual characters, and for species with male parental care as compared to those without, suggest that parasite resistance may be a general determinant of parasite-mediated sexual selection.     

The cause of parasitic infection in natural populations of Daphnia (Crustacea: Cladocera): the role of host genetics.

Disease patterns in nature may be determined by genetic variation for resistance or by factors, genetic or environmental, which influence the host-parasite encounter rate. Elucidating the cause of natural infection patterns has been a major pursuit of parasitologists, but it also matters for evolutionary biologists because host resistance genes must influence the expression of disease if parasite-mediated selection is to occur. We used a model system in order to disentangle the strict genetic component from other causes of infection in the wild. Using the crustacean Daphnia magna and its sterilizing bacterial parasite Pasteuria ramosa, we tested whether genetic variation for resistance, as determined under controlled conditions, accounted for the distribution of infections within natural populations. Specifically, we compared whether the clonally produced great-granddaughters of those individuals that were infected in field samples (but were subsequently 'cured' with antibiotics) were more susceptible than were the great-granddaughters of those individuals that were healthy in field samples. High doses of parasite spores led to increased infection in all four study populations, indicating the importance of encounter rate. Host genetics appeared to be irrelevant to natural infection patterns in one population. However, in three other populations hosts that were healthy in the field had greater genetic-based resistance than hosts that were infected in the field, unambiguously showing the effect of host genetic factors on the expression of disease in the wild.     

Sex-specific response of a mosquito to parasites and crowding

Host-parasite interactions are significantly influenced by the sex of the host and the environment in which the host is found. Sex-specific responses to parasite infection, however, may change according to the host environment. I examine the combined effect of parasite infection and crowding on males and females of the mosquito Aedes albopictus. At a high larval density, infected males experienced a greater relative reduction in body size than did infected females, whereas the pattern was reversed at low density. This experiment demonstrates the importance of the environment on sex-specific responses to parasites and contributes to a growing body of work examining sources of variation in host-parasite interactions.     

A genetic analysis of early development in pink (Oncorhynchus gorbuscha) and chum salmon (Oncorhynchus keta) at three different temperatures

A factorial mating design was employed in which five males were mated to each of five females in each of two stocks for both pink and chum salmon. The resulting embryos and alevins were incubated at constant water temperatures of 4, 8, and 16 degrees C for pink salmon and 3, 8, and 15 degrees C for chum salmon. Variation among families in alevin and fry survival rates, hatching, button-up time, length, and weight was the least at 8 degrees C. Heritability of traits directly correlated with fitness, such as survival rates and button-up time, was low at all temperatures (h2 less than or equal to 0.25). Maternal effects could account for a substantial portion of the variation in alevin and fry size characters. Nonadditive genetic variance accounted for more of the variation in fry size characters than in those of alevins. Negative genetic correlations were observed between embryo survival and subsequent alevin size and between hatching time and subsequent alevin and fry size. Genotype-temperature interactions could underlie a substantial amount of phenotypic variation in the developmental characters examined for both species.     

镜湖萼花臂尾轮虫种群内不同基因型克隆的生活史特征

应用生命表统计学等方法,比较了15、20、25和30℃4个温度下镜湖萼花臂尾轮虫种群内13个不同基因型克隆的生命表统计学参数和后代混交雌体百分率等生活史特征．结果表明：轮虫的世代时间、平均寿命、出生时的生命期望值、内禀增长率、净生殖率和后代混交雌体百分率对升高温度的反应均因克隆的不同而存在差异;温度、克隆及两者间的交互作用对其也都具有极显著影响．表明水环境在温度等方面所具有的时间异质性可能是镜湖萼花臂尾轮虫种群丰富的遗传变异得以维持的原因之一;自然选择出现在共存的克隆中,但其强度较低或作用时间较短是克隆共存的重要原因;遗传漂变可能在种群的基因型频率变化中起重要作用．不同基因型轮虫克隆的存在对其种群在水环境中的持续存在具有重要意义．     

Major histocompatibility complex heterozygote advantage and widespread bacterial infections in populations of Chinook salmon (Oncorhynchus tshawytscha)

Despite growing evidence for parasite-mediated selection on the vertebrate major histocompatibility complex (MHC), little is known about variation in the bacterial parasite community within and among host populations or its influence on MHC evolution. In this study, we characterize variation in the parasitic bacterial community associated with Chinook salmon ( Oncorhynchus tshawytscha ) fry in five populations in British Columbia (BC), Canada across 2 years, and examine whether bacterial infections are a potential source of selection on the MHC. We found an unprecedented diversity of bacteria infecting fry with a total of 55 unique bacteria identified. Bacterial infection rates varied from 9% to 29% among populations and there was a significant isolation by distance relationship in bacterial community phylogenetic similarity across the populations. Spatial variation in the frequency of infections and in the phylogenetic similarity of bacterial communities may result in differential parasite-mediated selection at the MHC across populations. Across all populations, we found evidence of a heterozygote advantage at the MHC class II, which may be a source of balancing selection on this locus. Interestingly, a co-inertia analysis indicated only susceptibility associations between a few of the MHC class I and II alleles and specific bacterial parasites; there was no evidence that any of the alleles provided resistance to the bacteria. Our results reveal a complex bacterial community infecting populations of a fish and underscore the importance of considering the role of multiple pathogens in the evolution of host adaptations.     

Rearing temperature influences flavivirus vector competence of mosquitoes

Culex annulirostris Skuse mosquitoes (Brisbane strain) were reared at 20 degrees C or 27 degrees C and the adult females were experimentally infected by feeding Murray Valley encephalitis virus (MVE). They were then maintained (a) in the insectary at 20 degrees C, after rearing at either 20 degrees C or 27 degrees C; (b) at ambient outdoor temperatures, range 12.2-28.9 degrees C, mean 19.6 degrees C; or (c) at 27 degrees C after rearing at 27 degrees C. There was no significant difference in rates of MVE infection or transmission when mosquitoes were reared and maintained constantly at 20 degrees C or 27 degrees C. However, for females kept at reduced temperature (i.e. mean = 19.6 degrees C or 20 degrees C after rearing at 27 degrees C), the infection and transmission rates of MVE were significantly reduced (2 x 8 replicates). This investigation illustrates that vector competence is depressed by decreasing temperatures for adult mosquitoes compared with those they experienced during development. Similar patterns were evident with previously published work on Japanese and St Louis encephalitis, dengue and yellow fever. The process appears to be reversible, i.e. increased temperature raises virus infection and transmission rates. It is concluded that, without incubation at warmer temperatures, flavivirus recovery from overwintering mosquitoes will be negatively biased.     

Effect of sexual segregation on host-parasite interaction: Model simulation for abomasal parasite dynamics in alpine ibex (Capraibex)

We investigated whether sexual segregation might affect parasite transmission and host dynamics, hypothesising that if males are the more heavily infected sex and more responsible for the transmission of parasite infections, female avoidance of males and the space they occupy could reduce infection rates. A mathematical model, simulating the interaction between abomasal parasites and a hypothetical alpine ibex (Capraibex) host population composed of its two sexes, was developed to predict the effect of different degrees of sexual segregation on parasite intensity and on host abundance. The results showed that when females tended to be segregated from males, and males were distributed randomly across space, the impact of parasites was the lowest, resulting in the highest host abundance, with each sex having the lowest parasite intensity. The predicted condition that minimises the impact of parasites in our model was the one closest to that observed in nature where females actively seek out the more segregated sites while males are less selective in their ranging behaviour. The overlapping of field observation with the predicted optimal strategy lends support to our idea that there might be a connection between parasite transmission and sexual segregation. Our simulations provide the biological boundaries of host-parasite interaction needed to determine a parasite-mediated effect on sexual segregation and a formalised null hypothesis against which to test future field experiments.     

Sexual selection explains more functional variation in the mammalian major histocompatibility complex than parasitism

Understanding drivers of genetic diversity at the major histocompatibility complex (MHC) is vitally important for predicting how vertebrate immune defence might respond to future selection pressures and for preserving immunogenetic diversity in declining populations. Parasite-mediated selection is believed to be the major selective force generating MHC polymorphism, and while MHC-based mating preferences also exist for multiple species including humans, the general importance of mate choice is debated. To investigate the contributions of parasitism and sexual selection in explaining among-species variation in MHC diversity, we applied comparative methods and meta-analysis across 112 mammal species, including carnivores, bats, primates, rodents and ungulates. We tested whether MHC diversity increased with parasite richness and relative testes size (as an indicator of the potential for mate choice), while controlling for phylogenetic autocorrelation, neutral mutation rate and confounding ecological variables. We found that MHC nucleotide diversity increased with parasite richness for bats and ungulates but decreased with parasite richness for carnivores. By contrast, nucleotide diversity increased with relative testes size for all taxa. This study provides support for both parasite-mediated and sexual selection in shaping functional MHC polymorphism across mammals, and importantly, suggests that sexual selection could have a more general role than previously thought.     

Effect of temperature on RNA synthesis in Escherichia coli CRT 266 (dna Bts) following infection with bacteriophage T3]

Infection of the temperature-sensitive E. coli CRT 266 (dnaBts) with T3-phages at the temperature of 30 degrees C and 35 degrees C, respectively, induced T3-specific RNA synthesis with a maximum rate at 7 min (30 degrees C) and 4.5 min (35 degrees C) after infection. At temperatures above 40 degrees C no T3-induced RNA synthesis could be observed. Infection of E. coli CR 34--45 (dnaB+) with T3 phages at 30 degrees C, 35 degrees C and at temperatures above 40 degrees C, however, produced T3-specific RNA synthesis. The maximum of T3-induced RNA synthesis could be observed between 7 min and 3 min depending on the temperature during infection. The inability to form T3-specific RNA after infection of E. coli CRT 266 at nonpermissive temperatures may be a cause for the absence of the formation of T3 phages and lysis of the host cells.     

EVOLUTION OF HOST-PARASITE DIVERSITY

Hosts and parasites often have extensive genetic diversity for resistance and virulence (host range). Qualitative diversity occurs when the success of attack is an all-or-nothing response that varies according to the genotypes of the host and parasite. Quantitative diversity occurs when the success of attack is a graded response that depends on additive genetic variation in the host and parasite. Community diversity occurs when parasites vary in the success with which they can attack different host species, leading to a mixture of specialists and generalists. I developed a series of models that classify components of host-parasite interactions according to whether they cause stabilizing or disruptive selection for resistance and virulence. Stabilizing selection reduces diversity by favoring a single optimal phenotype. Disruptive selection creates diversity by favoring a mixture of widely separated phenotypes. The evolution of maximal resistance and virulence are opposed by one of three forces: metabolic costs, frequency dependence, or negative genetic correlations among beneficial traits. The models predict that qualitatively inherited resistance and virulence traits typically cause greater diversity than quantitatively inherited traits. However, each natural system is composed of many stabilizing factors that reduce diversity and disruptive factors that promote diversity. I advocate a style of modeling in which families of related assumptions are compared by their equilibrium properties, and general conclusions from equilibrium properties are tested by complete dynamical analysis. The comparison among models highlights the need for empirical studies that compare levels of diversity among related host-parasite systems.