Wolbachia在山楂双叶螨中的感染及对寄主生殖的影响

【目的】共生菌Wolbachia在多种叶螨寄主中引起细胞质不亲和及适合度改变,影响寄主的生物学特性。山楂双叶螨Amphitetranychus viennensis是重要的果树害螨,常暴发成灾。本研究旨在明确Wolbachia在山楂双叶螨中的感染情况及对寄主生殖的影响。【方法】采集自然种群的山楂双叶螨,运用多位点序列分型技术（multilocus sequence typing, MLST）对其体内Wolbachia感染率及株系进行分析;通过杂交试验及生物学观察,分析感染Wolbachia对山楂双叶螨单雌产卵量、后代孵化率、性比及死亡率的影响。【结果】山楂双叶螨自然种群感染一种株系的Wolbachia （wVie）,该Wolbachia株系与小黑花椿象Orius strigicollis和丽蝇蛹集金小蜂Nasonia vitripennis中的Wolbachia株系亲缘关系较近,而与叶螨属Tetranychus叶螨感染的Wolbachia株系亲缘关系较远。Wolbachia与4种分化较小的线粒体单倍型相关联。Wolbachia感染雌虫与不感染雌虫产卵量没有显著差异（P>0.05）。不感染雌虫与感染雄虫交配,卵孵化率显著低于其他杂交组合 (P<0.05),但孵化率仍达近75%。各交配组合的后代性比及死亡率变化不明显（P>0.05）。【结论】Wolbachia在山楂双叶螨种群中的侵染历史较短,对山楂双叶螨的产卵力、后代性比、死亡率没有影响。Wolbachia在山楂双叶螨中诱导产生弱的CI表型。     

Host influences on reproduction and establishment of mouse-adapted Nippostrongylus brasiliensis (Nematoda)

Various influences of the host altered the establishment and reproduction of mouse-adapted Nippostrongylus brasiliensis. Establishment and egg production were greater in mature male than in female mice. Worm fecundity increased similarly in both sexes of mice prior to 41 days of host age. Male mice at 62 days of age or older harbored more N. brasiliensis than did females, but the number of nematodes was similar in younger hosts of both sexes. Crowding of male mice significantly increased the establishment of helminths, but not the production of eggs by N. brasiliensis. Pregnancy of the host significantly increased both establishment of and reproduction by N. brasiliensis. Ligation of the bile duct of the mouse reduced both establishment and fecundity of the nematode after subcutaneous or surgical infection with larvae and adults, respectively.     

Host location,survival and fecundity of the Oriental rat flea Xenopsylla cheopis (Siphonaptera: Pulicidae) in relation to black rat Rattus rattus (Rodentia: Muridae) host age and sex

Host choice and fecundity are two factors that may contribute to the variation in flea counts observed when assessing the potential risk of flea-borne transmission of pathogens from rodents to humans. Using the black rat, Rattus rattus Linnaeus, as host the effects of age and sex on host choice and fecundity of the Oriental rat flea, Xenopsylla cheopis Rothschild, were examined experimentally at 25 degrees C and 80% rh. During the first two days of emergence from cocoons, female fleas dominated the sex ratio by 4:1 but from the third day onwards this switched to a male-dominated sex ratio of 4:1. The sex of the flea did not influence their host-seeking behaviour. Newly emerged fleas of both sexes were not influenced by the rat's presence and at seven days old both sexes demonstrated similar levels of attraction toward the rat host. The sex of the rat did not affect flea host-seeking behaviour. There was a 50-70% decline in the initial number of adult fleas during the first week after their release onto a rat host, and this decline was greatest on juvenile rats. Flea fecundity was also significantly lower on juvenile rat hosts but no differences due to the sex of the rat were observed. This experimental study supports the hypothesis that differences in flea count due to host sex, reported in field surveys, result from sexual differences in host behaviour and not from discriminatory host-seeking behaviour by X. cheopis. Differences in flea count due to host age may be affected by differences in X. cheopis fecundity, which may itself be mediated by host behaviour such as grooming.     

Sex‐dependent infection causes nonadditive effects on kissing bug fecundity

The influence of parasites on host reproduction has been widely studied in natural and experimental conditions. Most studies, however, have evaluated the parasite impact on female hosts only, neglecting the contribution of males for host reproduction. This omission is unfortunate as sex‐dependent infection may have important implications for host–parasite associations. Here, we evaluate for the first time the independent and nonindependent effects of gender infection on host reproductive success using the kissing bug Mepraia spinolai and the protozoan Trypanosoma cruzi as model system. We set up four crossing treatments including the following: (1) both genders infected, (2) both genders uninfected, (3) males infected—females uninfected, and (4) males uninfected—females infected, using fecundity measures as response variables. Interactive effects of infection between sexes were prevalent. Uninfected females produced more and heavier eggs when crossed with uninfected than infected males. Uninfected males, in turn, sired more eggs and nymphs when crossed with uninfected than infected females. Unexpectedly, infected males sired more nymphs when crossed with infected than uninfected females. These results can be explained by the effect of parasitism on host body size. As infection reduced size in both genders, infection on one sex only creates body size mismatches and mating constraints that are not present in pairs with the same infection status. Our results indicate the fitness impact of parasitism was contingent on the infection status of genders and mediated by body size. As the fecundity impact of parasitism cannot be estimated independently for each gender, inferences based only on female host infection run the risk of providing biased estimates of parasite‐mediated impact on host reproduction.     

Effects of host age and host availability on developmental period,adult size,sex ratio,longevity and fecundity inLariophagus distinguendus Förster (Hymenoptera: Pteromalidae)

The effects of host age on parasitoid reproductive capacity are studied using the pteromalid parasitoid Lariophagus distinguendusFörster and its bruchid hosts, Callosobruchus chinensis (L.) and C. maculatus (F.). A series of experiments were performed to investigate relationships between age and size of host parasitized and the developmental period of pre-imaginal progeny, sex ratio, female size, longevity, fecundity and oviposition rate. There was no effect of host size on preimaginal parasitoid developmental period. Sex ratio varied from less than 5% females from young (small) hosts to 60% females from mature (large) hosts. Adult size, female longevity, fecundity, and oviposition rate were also positively related to host age. Females provided mature hosts lived longer than those provided either young hosts or no hosts, possibly because of an increased ability to host-feed from the larger hosts. The implications of these findings to parasitoid population reproductive capacity and host-parasitoid synchrony are discussed.     

Parasitism of Tuta absoluta (Lepidoptera, Gelechiidae) by Pseudapanteles dignus (Hymenoptera, Braconidae) under laboratory conditions

Laboratory studies were conducted to measure selected life history traits and the functional response of the parasitoid Pseudapanteles dignus (Muesebeck), a major enemy of Tuta absoluta (Meyrick) in tomato crops in South America. Newly mated P. dignus females were individually exposed to 10 host larvae in mines for 24 h. We determined developmental time from egg to pupal formation and pupal stage duration, female adult life span, fecundity, reproductive period, daily parasitism rate, and sex ratio of offspring. For the functional response experiment, treatments consisted of six host densities: 3, 5, 7, 10, 15, 20, or 30 larvae. The number and proportion of parasitized hosts were calculated for each density. The shape of the functional response curve was analyzed by logistic regression. P. dignus females attacked hosts daily, exhibiting modest lifetime fecundity (approximately 32 parasitized hosts per female) and a female-biased offspring sex ratio. Female adult life span was 36 d. P. dignus showed a type I functional response within the range of host densities tested. We observed that females detect and parasitize the host within a wide range of densities, including low densities. The functional response curve reached an asymptote at a mean density of six hosts per day and seemed not to be egg-limited. Percent parasitism was approximately 30%. The ecological implications of the results in relation to the potential of P. dignus for the biological control of T. absoluta in tomato are discussed.     

Factors affecting sex ratio in rearing ofCoeloides sordidator (Hym.: Braconidae)

Five factors known to affect the sex ratio (% of males) in parasitic Hymenoptera were investigated forCoeloides sordidator, a parasitoid ofPissodes weevils. The host age, the age of ovipositing females, and the host of origin had a significant impact on the sex ratio of offspring. In contrast, the number of ovipositing females had an insignificant effect on sex ratio whereas the effect of host density could not be clearly defined. The sex ratio decreased with host age, probably because, like many other hymenopteran parasitoids, females tend to lay male eggs on small hosts and female eggs on larger hosts in order to maximize the size and fitness of their female offspring. The sex ratio also varied with the age of the mother, younger females laying more male eggs and older females more female eggs. The host of origin also had an influence on sex ratio. The strain fromPissodes castaneus was significantly more male-biased than the strain fromP. validirostris, which corroborates previous observations made on field populations     

Impact of mating on Anagyrus kamali Moursi (Hym., Encyrtidae) lifetime fecundity, reproductive longevity, progeny emergence and sex ratio

Abstract: The solitary endoparasitoid Anagyrus kamali Moursi (Hym., Encyrtidae) and the Hibiscus mealybug Maconellicoccus hirsutus Green (Hom., Pseudococcidae), were used as a parasite/host model to test the effect of mating on several fitness parameters, i.e. longevity, lifetime fecundity, progeny emergence and sex ratio. At 27 ± 2°C, 8 h light : 16 h dark, mating significantly affected the survival of male parasitoids. Virgin males lived longer (32.2 ± 9.51 days) than mated males (23.9 ± 7.52 days). Female longevity (40.7 ± 16.3 days for virgins and 36.2 ± 10.7 days for mated females) was not affected by mating. The lifetime fecundity of female parasitoids and their oviposition period was not significantly affected by mating. However, the number of hosts parasitized was greater for mated wasps (7.54 ± 4.85 hosts parasitized/day) compared with virgin ones (5.12 ± 2.19 hosts parasitized/day). This resulted in greater progeny production from mated A. kamali females. The progeny of virgin females consisted only of males, whereas the mated ones had a more female‐biased sex ratio. The lowest sex ratio (0.41 M/F ± 0.123) was attained when females had free access to males and were multi‐mated.     

Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size

Adaptiveness of sex ratio control by the solitary parasitoid wasp Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size was studied, by examining whether differential effects of host size on the fitness of resulting wasps are to be found between males and females. The offspring sex ratio (male ratio) decreased with increasing host size. Larger hosts yielded larger wasps. Male larvae were less efficient in consuming larger hosts than female larvae. No significant interaction in development time was found between parasitoid sex and host size. Larger female wasps lived longer than smaller females, while longevity of male wasps did not increase with increasing wasp size. Smaller males were able to mate either with small or with large females, while larger males failed to mate with small females. Larger female wasps had a greater number of ovarioles and mature eggs at any one time than smaller females, although the number of eggs produced per host-feeding was not influenced by female wasps. Thus, the differential effect of host size on the fitness of males and females exists in I. naranyae. The basic assumption of the host-size model was therefore satisfied, demonstrating that sex ratio control by I. naranyae in response to host size is adaptive.     

Transmission pattern and intraspecific competition as determinants of population structure in pinworms (Oxyurida: Nematoda).

The transmission pattern of Zonothrix columbianus (Nematoda: Oxyurida) in its host Tropisternus columbianus (Coleoptera: Hydrophilidae), an aquatic beetle, was studied to determine whether parasites were dispersed with their hosts and to examine the possible role of intraspecific competition in limiting population size. Beetles were sampled at regular intervals from fall 1986 through fall 1989 and examined for worms. Worms, absent in larval stages of the host, were uncommon in newly metamorphosed beetles and therefore probably do not infect adult stages until after they have dispersed; worms are not dispersed with the host. Prevalence reached its lowest points in spring and fall when newly metamorphosed beetles were most common, but it was near 100% for most of the year. Worms were uniformly distributed in the host population. Many hosts had exactly 1 male and 1 female worm; the high prevalence suggests that this infrahost population results from interference competition between males on the one hand and females on the other. Only 3 of 285 beetles contained more than 1 male. Females shared the host with members of the same sex more commonly than males, but females from hosts harboring more than 1 female had significantly fewer eggs than lone females in hosts. Numbers of adult stages of beetles were estimated during spring, summer, and fall of 1989 and were lower in early spring and late fall. Because worms do not disperse with hosts, the panmictic unit could be estimated from the number of infected beetles; this probably was about 50 individuals during the winter bottleneck.     

Infection of Harmonia axyridis (Coleoptera: Coccinellidae) by Hesperomyces virescens (Ascomycetes: Laboulbeniales): role of mating status and aggregation behavior

The ectoparasitic fungus Hesperomyces virescens was studied on Harmonia axyridis in North Carolina, in the southeastern United States. A primary goal was to investigate transmission of the disease by examining the correlation between the pattern of fungal infection and seasonal change in host behavior. Beetles were collected as they arrived at their winter quarters at two sites; in one site they were also subsampled at mid- and late winter. Insects were sexed and weighed, fungal thalli were counted, and their location on the host body mapped; spermathecae of females were examined for sperm. Infection levels varied between sites, differed significantly between the sexes in one site but not the other, and increased by approximately 40% during winter. The distribution of thalli on the body changed seasonally, in concert with behavioral changes in the host. At fall flight, thalli were found most often on the posterior elytra of mated females, virgin females, and males. This is suggestive that the disease had been spread among both sexes via successful and failed copulation attempts; however, the relatively low incidence of infection on the male venter does not fit the sexual transmission scenario. During winter, thallus location shifts in concert with beetle aggregation behavior, with infections more often located on the head and legs. Fresh weight of beetles decreased by approx. 20% during winter, but was not affected by disease status. Prior to spring flight, uninfected females were preferred as mating partners, but the probable relationship between female age and infection status complicates interpretation of the data.     

Sequential and concurrent exposure of flour beetles ( Tribolium confusum ) to tapeworms ( Hymenolepis diminuta ) and pesticide (diatomaceous earth)

The response of Tribolium confusum to sublethal levels of 2 environmental stressors was studied, i.e., parasitic infection represented by the cestode Hymenolepis diminuta , and a physical stressor represented by the natural pesticide diatomaceous earth (DE). These were applied sequentially (DE, then infection) to detect indirect or carryover effects of DE, and concurrently (DE applied immediately after exposure to parasites and DE presence maintained throughout the infection) to detect direct effects of DE. DE alone, but not parasitism alone, produced significant host mortality, and concurrent treatment with DE and parasitism did not increase mortality over DE alone. Parasite abundance was significantly higher following sequential, but not concurrent, DE exposure. Parasite abundance in mated hosts was significantly higher than in virgin hosts. Parasitic infection resulted in significantly fewer eggs retained in the oviduct of beetles, but there was no difference in the number of eggs that accumulated in the culture medium and no difference in the surface-seeking behavior of beetles. Mating status of beetles in all treatments, and DE exposure in concurrent treatments significantly increased their surface-seeking behavior. Concurrent exposure to DE also resulted in a 4- to 6-fold increase in host egg numbers that accumulated in the culture medium. Although DE exposure increased parasite numbers in the beetles, these 2 stressors otherwise appeared to act independently.     

Sex ratios in field populations of two parasitoids (Hymenoptera: Chalcidoidea) of Coccus hesperidum L. (Homoptera: Coccidae)

We tested several assumptions and predictions of host-quality-dependent sex allocation theory (Charnov et al. 1981) with data obtained for the parasitoid Metaphycus stanleyi Compere on its host, brown soft scale (Coccus hesperidum L.), in a California citrus grove and in the laboratory. Scales ceased growing after parasitization by M.?stanleyi. Thus, M. stanleyi may gauge host quality (=size) at oviposition. Host size positively influenced adult parasitoid size, and parasitoid size in turn influenced adult longevity of M. stanleyi. However, parasitoid fitness gains with host size and adult size were similar in males versus females. Sex allocation to individual hosts by M. stanleyi depended on host size; females consistently emerged from larger hosts than males. Host size was important in a relative sense; the mean host sizes of females versus males, and of solitary versus gregarious parasitoids varied with the available host size distribution. The offspring sex ratio of M. stanleyi reflected the available host size distribution; the sex ratio of emerging parasitoids varied with the available host size distribution. We did not detect a “critical host size” below which males emerged, and above which females emerged; rather, only females emerged from hosts in the upper size range, and a variable ratio of males and females emerged from hosts in the lower size range. We conclude that the sex ratio of field populations of M.?stanleyi is driven largely by the available size distribution of C. hesperidum. In addition, we tested predictions resulting from theoretical analyses of sex allocation in autoparasitoids with data obtained on Coccophagus semicircularis (Förster) parasitizing brown soft scale in the field. The sex ratio of C. semicircularis was consistently and strongly female biased (ca. 90% females). Based on available theoretical analyses, we suggest that this sex ratio pattern may have resulted from a very low encounter rate of secondary hosts coupled with a strong time limitation in C. semicircularis females. This explanation was the most plausible given constraints stemming from the detection of secondary hosts, their variable location within primary hosts, and their handling times. Finally, the size of hosts which yielded single versus multiple parasitoids, and the sizes of these parasitoids, were compared. These comparisons suggested that: (1) M. stanleyi females gauge host sizes precisely, and in terms of female offspring; thus a fitness penalty is not incurred by females which share a host, while males benefit from sharing a host, and; (2) instances where multiple C. semicircularis emerged from a single host were probably the result of parasitism by different females, or during different encounters by a single female.     

Role of Volatile Semiochemicals in the Host and Mate Location Behavior of Mallodon dasystomus (Coleoptera: Cerambycidae)

Little is known of the role semiochemicals play in the mating systems of longhorned beetles (Coleoptera: Cerambycidae) in the primitive subfamily Prioninae. Mallodon dasystomus (Say), the hardwood stump borer, is a widely distributed prionine native to the southern US. Preferred hosts of M. dasystomus include oak, sweetgum, sugarberry and hackberry; although they also colonize a variety of other hardwoods. Here, we study the mate location behavior of M. dasystomus by testing the hypotheses that the sexes are mutually attracted to volatiles emanating from the larval host and that females release a volatile pheromone that is attractive to males alone. In a Y-tube olfactometer, male and female M. dasystomus responded to volatiles from host material (i.e., sweetgum and sugarberry). However, only males responded to females in the olfactometer, suggesting that females release a volatile sex pheromone. In choice experiments conducted in a greenhouse, we determined that both males and females prefer host over non-host material. In further bioassays in the greenhouse, males chose host material containing a live female over that containing a live male or host material alone. These findings are further evidence of the critical role host volatiles and pheromones play in mating systems of longhorned beetles.     

The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Pteromalus puparum is a gregarious parasitoid of many butterfly pupae. Adult size, mortality, and sex ratio of P. puparum, as a parasitoid of Papilio xuthus, were unit weight of the host. Effects of female size on fecundity, wing load, and longevity were also examined.The highest total weight of progeny from the host was attained when the number of eggs per gram of the host was approximately 150. Positive correlations were observed between the size of the females and their fecundity and wing load. The maximum longevity of the female kept with honey but without hosts was attained when the initial number of parasitoids per g of the host was 150.Considering the total fecundity of all female progeny, the reproductively most efficient number of eggs to be deposited per g of the host was estimated to be approximately 300. However, as shortage of food for the adult females strongly affects their fecundity, the reproductively most efficient number of eggs to be deposited per g of the host was about 70 when the adult female progeny was not provided with food.The optimal number of eggs to be deposited when the emale oviposits in the host under field conditions is discussed.     

Some factors affecting reproduction and sex-ratio of the predacious mite Amblyseius deleoni (Muma and Denmark) (Acari: Phytoseiidae)

Abstract

Several factors were examined to determine their effect on the reproduction and sex ratio in the predacious mite Amblyseius deleoni (Muma and Denmark), in the laboratory. The factors investigated included multiple matings, duration of copulation, capacity of male for mating in excess of females and age of mating females and males. The factors included also, the host plant leaf texture, food deprivation during immature and adult stages, and prey (Tetranychus urticae Koch) density. The results indicated that females of A. deleoni require multiple matings to maximize their reproductive potential, also when copulation was allowed for increasing periods of time, there was a gradual increase in total egg production and oviposition period. A male showed a high reproductive ability for more than 15 days and was able to mate more than once in excess of females. Age of females has an influence on fecundity and sex ratio; old females decreased egg production and produced proportionally more male progeny compared with young females. Similarly, the highest number of eggs deposited per female A. deleoni was reported, when female mated with a young male (0-day old). In addition, males of A. deleoni (at any age) were able to inseminate the females. Results from host plant leaf texture indicated that guava leaf gave the highest reproduction rate, while the fig leaf gave the least female fecundity. Neither the reproductive rate nor the sex ratio of the progeny of females crossed by normal or experimental males had been influenced by the food deprivation during immature stages. A significant lower fecundity was recorded on female's A. deleoni when exposed to different food deprivation programmes during adult stage. The number of eggs laid by the predator female increased with increasing prey density of T. urtice to a maximum of 2.04 eggs deposited per day at a prey density of 30 protonymphs of T. urticae as a prey. As the level of prey density was increased, there was a shift in sex ratio towards an increased proportion of females.     

Occurrence of the internal parasite Portunion sp. (Isopoda: Entoniscidae) and its effect on reproduction in intertidal crabs (Decapoda: Grapsidae) from New Zealand

The parasite fauna of 4 intertidal grapsid crabs from New Zealand was studied between 1998 and 2000. The occurrence of an undescribed entoniscid species, Portunion sp. (Isopoda: Epicaridea), is presented. Portunion sp. was found in Cyclograpsus lavauxi (34.1%, n = 1650), Hemigrapsus crenulatus (19.0%, n = 2300), and Helice crassa (11.6%, n = 825) but was absent from Hemigrapsus sexdentatus (n = 636). Parasitized hosts contained mostly 1 female Portunion sp. but occasionally up to 7 females were found. One to 3 dwarf males typically occurred on each mature female Portunion sp. Most developmental stages of female Portunion sp. were found throughout the year, demonstrating that reproduction and infection occurred continuously. Portunion sp. differentially affected male and female hosts. Prevalence was generally higher in male hosts than in female hosts and increased significantly with host size. Female hosts were castrated, whereas males were not. Portunion sp., therefore, influenced the operational sex ratio in its host species, causing a more male-biased ratio.     

Role of host volatiles in mate location by the Japanese pine sawyer, Monochamus alternatus hope (Coleoptera: Cerambycidae)

We evaluated the responses of male and female Monochamus alternatus Hope (Coleoptera: Cerambycidae) to various terpenes commonly associated with host trees. Electroantennogram (EAG) tests were conducted with 12 plant volatile compounds and ethanol. Antennae of both sexes were highly sensitive to (R)-(+)-alpha-pinene, (+)-3-carene, (-)-beta-pinene, and terpinolene. Both sexes of M. alternatus were attracted by traps baited with (+)-alpha-pinene, (-)-beta-pinene, (+)-3-carene, or terpinolene. Our results support the first of the three-stage hypothesis posed by Ginzel and Hanks that suggests that location of stressed trees by cerambycids involves three stages: (1) both sexes locate larval hosts by using plant volatiles as kairomones; (2) males produces sex pheromones to attract females after both sexes land on the larval hosts; (3) males and female recognize each other by contract pheromones in their epicuticular wax layer. Males and females showed differences in their EAG responses to several compounds, including (R)-(+)-alpha-pinene, (-)-beta-pinene, myrcene, (+)-3-carene, (R)-(+)-limonene, terpinolene, and trans-caryophyllene. In all cases, males exhibited greater sensitivity than females. In laboratory assays, male M. alternatus showed strong preference for 1% (+)-alpha-pinene and 1% (-)-beta-pinene over other compounds. In field assays, traps baited with (+)-alpha-pinene, (-)-beta-pinene, (+)-3-carene, or terpinolene caught more beetles than control traps. We found strong male bias in beetle catches in baited traps and those captured on the stem of stressed trees despite a strong female bias in emerging beetles in 2004. We hypothesize that male M. alternatus are more responsive than females to plant volatiles and that males have more capacity than females in finding mating locations.     

The relationships between host selection and subsequent larval performance in three free-living graminivorous sawflies

1. Selection of host plants for oviposition by females of three graminivorous sawfly species, Dolerus puncticollis C. G. Thomson, Dolerus picipes Klug. and Pachynematus clitellatus Lepeletier, was investigated experimentally using three grass species, Lolium perenne L., Festuca rubra L. and wheat, Triticum aestivum L. cv. Hunter.
2. Both species of Dolerus showed significant preferences for one grass over the other two, D. puncticollis preferring L. perenne and D. picipes selecting F. rubra. P. clitellatus females used L. perenne and F. rubra equally for oviposition, but avoided wheat.
3. The results correlated closely with larval performance trials, with D. puncticollis larvae having maximal survival and growth rates on L. perenne , and D. picipes larval performance being maximized on F. rubra. P. clitellatus larvae survived well on both grasses, but had a high level of mortality on wheat.
4. P. clitellatus larval growth rates were significantly higher on L. perenne than on F . rubra . Individual P. clitellatus adult females developing from larvae reared on L. perenne reached a significantly greater dry weight than those reared on F. rubra , but there was no significant difference in weight between males reared on the two hosts. There was some evidence that this disproportionate effect of host plant quality on the weight gain of the two sexes was translated into a bias in sex ratio of the eggs laid towards a greater proportion of females on L. perenne than on F. rubra . This manipulation of sex ratio would have a potential benefit for ovipositing females through disproportionate fecundity gains for female offspring.     

Virulence is context-dependent in a vertically transmitted aquatic host–microparasite system

Recent work has suggested that the outcomes of host–symbiont interactions can shift between positive, neutral and negative depending on both biotic and abiotic conditions. Even organisms traditionally defined as parasites can have positive effects on hosts under some conditions. For a given host–parasite system, the effects of infection on host fitness can depend on host vigour, route of transmission and environmental conditions. We monitored sublethal microsporidian infections in populations of Gammarus pseudolimnaeus (Amphipoda: Gammaridae) from four cool water streams in southwestern Michigan, USA. Our objectives were to: (i) infer the mechanism of transmission (horizontal, vertical or mixed) from observed effects of infection on host fitness, (ii) determine if the magnitude of the effects on host fitness is a function of parasite load (infection intensity) compared with simple presence or absence of infection, and (iii) determine if there is variation in parasite effects on host fitness in isolated populations. PCR and DNA sequence analyses revealed that there were two microsporidia present among the four host populations: Dictyocoela sp. and Microsporidium sp. PCR screening of a subset of infected hosts showed that Dictyocoela sp. accounted for 90% of infections and was present in all four G. pseudolimnaeus populations, while Microsporidium sp. was found in two populations but was only relatively common in one. We found very low prevalence in males (∼5%), but high prevalence in females (range: 37–85%). Female fitness was positively associated with infection in two streams, resulting from either higher fecundity or more reproductive bouts. Infection had a negative effect on the number of reproductive bouts in a third population, and no effect on fecundity in a fourth population. Infection intensity explained additional variation in fecundity in one population; females with intermediate infection intensity had higher fecundity than females with either light or heavy infection intensity. Given the high prevalence of infection in females compared with males and the generally weak negative fitness effects coupled with some positive fitness effects, it is likely that both Dictyocoela sp. and Microsporidium sp. are primarily vertically transmitted, feminizing microsporidia. Our results suggest that microsporidian effects on G. pseudolimnaeus fitness were context-dependent and varied with host sex and local environment.