Sexual dimorphism in immune function changes during the annual cycle in house sparrows

Difference between sexes in parasitism is a common phenomenon among birds, which may be related to differences between males and females in their investment into immune functions or as a consequence of differential exposure to parasites. Because life-history strategies change sex specifically during the annual cycle, immunological responses of the host aiming to reduce the impact of parasites may be sexually dimorphic. Despite the great complexity of the immune system, studies on immunoecology generally characterise the immune status through a few variables, often overlooking potentially important seasonal and gender effects. However, because of the differences in physiological and defence mechanisms among different arms of the immune system, we expect divergent responses of immune components to environmental seasonality. In male and female house sparrows (Passer domesticus), we measured the major components of the immune system (innate, acquired, cellular and humoral) during four important life-history stages across the year: (1) mating, (2) breeding, (3) moulting and (4) during the winter capture and also following introduction to captivity in aviary. Different individuals were sampled from the same population during the four life cycle stages. We found that three out of eight immune variables showed a significant life cycle stage × sex interaction. The difference in immune response between the sexes was significant in five immune variables during the mating stage, when females had consistently stronger immune function than males, while variables varied generally non-significantly with sex during the remaining three life cycle stages. Our results show that the immune system is highly variable between life cycle stages and sexes, highlighting the potential fine tuning of the immune system to specific physiological states and environmental conditions.     

Variation in sex pheromone emission does not reflect immunocompetence but affects attractiveness of male burying beetles—a combination of laboratory and field experiments

Life history theory predicts a trade-off between male sexual trait expression and immunocompetence. Using burying beetles, Nicrophorus vespilloides, as a model, we investigated the relationship between male immune function, sex pheromone emission, and attractiveness under field conditions. In the first experiment, we tested whether there is a positive correlation between immune capacity, sex pheromone characteristics (quantity, relative composition, and time invested in pheromone emission), and male attractiveness. As a measurement of immune capacity, we used an individual’s encapsulation ability against a novel antigen. In the second experiment, we specifically examined whether a trade-off between chemical trait expression and immune function existed. To this end, we challenged the immune system and measured the subsequent investment in sex pheromone emission and the attractiveness of the male under field conditions. We found that a male’s immunocompetence was neither related to the emission of the male’s sex pheromone nor to its attractiveness in the field. Furthermore, none of the immune-challenge treatments affected the subsequent investment in pheromone emission or number of females attracted. However, we showed that the same males that emitted a high quantity of their sex pheromone in the laboratory were able to attract more females in the field. Our data suggest that the chemical signal is not a reliable predictor of a male’s immunocompetence but rather is a general important fitness-related trait, with a higher emission of the sex pheromone measured in the laboratory directly affecting the attractiveness of a male under field conditions.     

Hunting behaviour and breeding performance of northern goshawks Accipiter gentilis, in relation to resource availability, sex, age and morphology

Animal territories that differ in the availability of food resources will require (all other things being equal) different levels of effort for successful reproduction. As a consequence, breeding performance may become most strongly dependent on factors that affect individual foraging where resources are poor. We investigated potential links between foraging behaviour, reproductive performance and morphology in a goshawk Accipiter gentilis population, which experienced markedly different resource levels in two different parts of the study area (rabbit-rich vs. rabbit-poor areas). Our analyses revealed (1) that rabbit abundance positively affected male reproductive output; (2) that age, size and rabbit abundance (during winter) positively affected different components of female reproductive output; (3) that foraging movements were inversely affected by rabbit abundance for both sexes (for females, this may mainly have reflected poor provisioning by males in the rabbit-poor area); (4) that younger breeders (both in males and females) tended to move over larger distances than older individuals (which may have reflected both a lack of hunting experience and mate searching); and (5) that male body size (wing length) showed some covariation with resource conditions (suggesting possible adaptations to hunting agile avian prey in the rabbit-poor area). Although we are unable to establish firm causal relationships with our observational data set, our results provide an example of how territory quality (here, food abundance) and individual features (here, age and morphology) may combine to shape a predator's foraging behaviour and, ultimately, its breeding performance.     

Form and nature of precopulatory sexual selection in both sexes of a moth

Sexual selection is a process that operates through intrasexual competition and intersexual choice for reproduction in both sexes. Here, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), aiming to infer the form and nature of precopulatory sexual selection in males and females. We show that, although a number of traits measured in each sex are correlated with mating success, the primary selection trait in females appears to be abdominal thickness and that in males is aedeagus length. As the female’s abdominal thickness is a reliable signal about the number and developmental stage of eggs, males who select females with thicker abdomens for mating will gain reproductive benefit, i.e., fertilizing more mature eggs. For females, earlier maturation of their eggs makes the females more likely to achieve mating earlier in an uncertain world where there is no guarantee that they will find more males in the future. Sexual selection appears to be the important force behind the evolution of fast egg maturation in females. We show that, under a male-biased sex ratio, more than 20% of mating fails within a few minutes after the aedeagus has penetrated into the female’s genitalia, suggesting that females can assess the features of the male aedeagus before allowing insemination to occur. Dissection and examination of both sexes suggest that a longer aedeagus enhances mating and fertilization efficiency in this species, supporting the notion that sexual selection is a primary force in the evolution of genital variance.     

Colour change in a structural ornament is related to individual quality,parasites and mating patterns in the blue tit

Carry-over effects refer to processes that occur in one season and influence fitness in the following. In birds, two costly activities, namely reproduction and moult, are restricted to a small time window, and sometimes overlap. Thus, colour in newly moulted feathers is likely to be affected by the costs of reproduction. Using models of bird vision we investigated male colour change in a free-living population of blue tits (Cyanistes caeruleus) in three sampling occasions: spring 1, winter and spring 2. We related crown, tail, breast and cheek feather colouration after the moult (winter) to the intensity of infections by blood parasites during reproduction (spring 1). In the following spring (spring 2), we explored mating patterns with respect to changes in feather colour (springs 1 vs. 2). Males that were less intensely infected by the malaria parasite Plasmodium while breeding showed purer white cheek feathers in winter, which may indicate higher feather quality. Increased brightness in the white cheek was associated with better body condition during reproduction. In the following season, males with brighter cheeks paired with females that had noticeably brighter cheek patches compared to the male’s previous mate. These results suggest that the conditions experienced during reproduction are likely to affect moult and thus feather colouration, at least in the white patch. High quality individuals may allocate resources efficiently during reproduction increasing future reproductive success through variation in mating patterns. Carry-over effects from reproduction might extend not only to the non-breeding phase, but also to the following breeding season.     

Sex- and age-related differences in the timing and body condition of migrating Reed Warblers Acrocephalus scirpaceus and Sedge Warblers Acrocephalus schoenobaenus

The migration strategies of birds may vary strongly between species and also between age and/or sex groups. We studied the autumn migration and body condition of molecularly sexed Reed Warblers Acrocephalus scirpaceus and Sedge Warblers Acrocephalus schoenobaenus (211 and 208 ind., respectively) at a stopover site on Lake Druzno, Northern Poland, in 2008. Immature male Reed Warblers were caught significantly later than females (median dates 9 days later), but in the Sedge Warbler, both sexes of immatures migrated at about the same time. Adult males and females of both species did not differ in their time of migration. Adult and immature males of both species were larger (wing length and body mass) than females. In both species, fat reserves were similar in both sexes of both age classes. Adults of both sexes of Reed and Sedge Warbler were generally caught earlier than immatures. In both species, the body mass and fat reserves of immatures were generally less than in adults. The autumn protogyny of immature Reed Warblers may allow smaller females to limit competition with bigger males during migration and at the wintering grounds. In the Sedge Warbler, which tends to match its migration to peak of occurrence of superabundant food at stopover sites, both sexes gain an advantage from migrating at the same time. Since part of the measured wing length variation in both species was explained by sex differences, temporal trends in wing length recorded at stopover sites should be interpreted with caution.     

Temporal variation in size-assortative mating and male mate choice in a spider with amphisexual care

Males should be more selective when they have a high investment in reproduction, especially in species with biparental or paternal care. In this context, male mate choice can promote size-assortative mating (SAM) when (1) large males win intrasexual disputes, (2) large females are more fecund, and (3) males prefer larger females to smaller ones. In the spider Manogea porracea, males exhibit high reproductive investment by building their webs above those of females and exhibiting extended care of offspring in the absence of females. Under these circumstances, we expect the occurrence of SAM and male preference for large females. Herein, we performed observations and experiments in the field to evaluate the hypotheses that (1) M. porracea mates assortatively by size and (2) SAM is influenced by male mate choice. Furthermore, we measured variables that could affect mating patterns, the sex ratios, and densities of both sexes. Pairing in M. porracea was positively size-assortative in 2012, but not in 2013. Large males won most disputes for mates and preferred larger females, which produced more eggs. The inconsistency in detection of SAM was due to population dynamics, namely variations in sex ratio and population density across the breeding season. Furthermore, we found that the significance of male mate choice on sexual selection of body size in M. porracea strongly depends on the competition intensity for mating opportunities. The traditional sexual selection hypothesis of SAM needs to be reviewed and must include measures of competition intensity.     

Variations of immune parameters in terrestrial isopods: a matter of gender, aging and Wolbachia

Ecological factors modulate animal immunocompetence and potentially shape the evolution of their immune systems. Not only environmental parameters impact on immunocompetence: Aging is one major cause of variability of immunocompetence between individuals, and sex-specific levels of immunocompetence have also been frequently described. Moreover, a growing core of data put in light that vertically transmitted symbionts can dramatically modulate the immunocompetence of their hosts. In this study, we addressed the influence of gender, age and the feminising endosymbiont Wolbachia (wVulC) on variations in haemocyte density, total PO activity and bacterial load in the haemolymph of the terrestrial isopod Armadillidium vulgare. This host–symbiont system is of particular interest to address this question since: (1) wVulC was previously shown as immunosuppressive in middle-aged females and (2) wVulC influences sex determination. We show that age, gender and Wolbachia modulate together immune parameters in A. vulgare. However, wVulC, which interacts with aging, appears to be the prominent factor interfering with both PO activity and haemocyte density. This interference with immune parameters is not the only aspect of wVulC virulence on its host, as reproduction and survival are also altered.     

Social Rank, Stress, Fitness, and Life Expectancy in Wild Rabbits

 Wild rabbits of the two sexes have separate linear rank orders, which are established and maintained by intensive fights. The social rank of individuals strongly influence their fitness: males and females that gain a high social rank, at least at the outset of their second breeding season, have a much higher lifetime fitness than subordinate individuals. This is because of two separate factors: a much higher fecundity and annual reproductive success and a 50% longer reproductive life span. These results are in contrast to the view in evolutionary biology that current reproduction can be increased only at the expense of future survival and/or fecundity. These concepts entail higher physiological costs in high-ranking mammals, which is not supported by our data: In wild rabbits the physiological costs of social positions are caused predominantly by differential psychosocial stress responses that are much lower in high-ranking than in low-ranking individuals. Received: 23 February 1999 / Accepted: 17 March 1999     

Sexual harassment and female gregariousness in the South American sea lion, Otaria flavescens

Most colonial pinnipeds form extreme clusters of breeding females that cannot be entirely explained by the distribution of sites for reproduction. Avoidance of male harassment has been postulated as an important determinant of reproductive aggregation in this group of mammals. Female gregariousness can reduce harassment by resident males by two mechanisms; directly by the 'dilution effect' or indirectly because resident males that defend large female groups are less harassing. In order to investigate the relationship between male harassment and female gregariousness in relation to the size of breeding groups, we analysed the behaviour of dominant males and their females in a breeding colony of Otaria flavescens. Females in large breeding groups received less harassment by resident males due to dilution effects and because males that defended a large group interacted less frequently with females than males with small groups.     

Carotenoid-dependent coloration of male American kestrels predicts ability to reduce parasitic infections

The signaling function of sexually selected traits, such as carotenoid-dependent avian plumage coloration, has received a great deal of recent attention especially with respect to parasitism and immunocompetence. We argue that parasite-mediated models of sexual selection may have an implicit temporal component that many researchers have ignored. For example, previous studies have demonstrated that carotenoid-dependent traits can signal past parasite exposure, current levels of parasitism, or the ability of individuals to manage parasitic infections in the future. We examined repeated measures of carotenoid-dependent skin color and blood parasitism in American kestrels (Falco sparverius) to distinguish whether coloration might signal current parasitism or the potential to deal with infections in the future. We found no evidence that coloration was related to current levels of parasitism in either sex. However, coloration of males significantly predicted their response to parasitism; males with bright orange coloration during prelaying, when mate choice is occurring, were more likely than dull yellow males to reduce their levels of infection by the time incubation began. Coloration during prelaying may advertise a male’s health later in the breeding season. For kestrels, the ability to predict future health would be highly beneficial given the male’s role in providing food to his mate and offspring. Coloration of females was not a significant predictor of parasitism in the future, and we provide several possible explanations for this result.     

Immune challenges and visual signalling in tree frogs

In animals, mate-choice is often based on sexual signals that carry information and help the receiver make the best choice to improve the receiver’s fitness. Orange visual sexual signals have been hypothesised to carry immune information because they are often due to carotenoid pigments which are also involved in immunity response. Although many studies have focused on the direct relationships between coloration and immunocompetence, few studies have simultaneously studied immunocompetent response and coloration variation after an immune challenge. We tested this hypothesis on starved and ad libitum-fed males of the European tree frog Hyla arborea. Our results show that male coloration is not a reliable indicator of its immune response capacity in this species. However, after an immune challenge induced by a PHA (Phaseolus vulgaris phytohaemagglutinin) injection, starved males presented a significant coloration loss and this alteration was related to the immune response intensity. Taken together, these results suggest that the brighter (lighter) coloration may be used as a cue by female to exclude males with a recent immune challenge, due to diseases or parasites for example.     

One trait, many signals: different information on male quality is enclosed within the same trait in a blenny fish

Colourful ornaments are traditionally evaluated as one trait. However, they could consist of several components, such as total size, colour intensity and extension, each possibly bearing its own message about one aspect of quality. Males of the blenny Salaria pavo exhibit a colourful head crest and solely care for eggs. During the breeding season, the head crest shows a yellow colouration, the intensity and relative extension of which are independent of crest size. Here, we show that: (1) carotenoids are responsible for the head crest yellow patch; (2) activating the immune system by injecting the bacterial antigen lipopolysaccharides affects both the intensity and extent of the yellow colouration; and (3) females assess males on the basis of colour patch expression. However, the response of the yellow patch to the immune challenge was dependent on head crest size. Indeed, males with a larger head crest reacted better to the simulated infection, sustaining a level of yellow patch close to pre-challenge size.     

Sexually dichromatic coloration reflects size and immunocompetence in female Spanish terrapins, Mauremys leprosa

Many studies have shown the importance of colorful ornamentation in mate choosiness or intrasexual conflict. However, research on color ornaments has focused mainly on birds, lizards or fish, but remains practically unknown in other animal groups such as turtles. In addition, female ornaments and their relation with sexual selection also remain almost unknown. Here, we measured the coloration of the shell and the limb stripes of male and female Spanish terrapins Mauremys leprosa and explored the existence of sexual dichromatism and the relation of color characteristics with body size and health state estimated from the immune response to the injection of an antigen (phytohaemagglutinin test). Our results showed that shell coloration, which could be constrained by natural selection to be cryptic, changed with body size, but did not differ between sexes. In contrast, females had brighter and less ultraviolet-saturated and more orange-saturated limb stripes than males. In females, interindividual variation in limb stripe coloration was related with body size and immune response suggesting that this coloration may inform honestly about multiple traits that could be important in sexual selection. In contrast, coloration of limb stripes of males was duller than in females, and was not related with any trait suggesting that coloration is not important in sexual selection for males.     

Mate guarding and territorial aggression vary with breeding synchrony in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Male paternity assurance behaviour during the female fertile period has been widely documented amongst birds. In contrast, how sex-specific behavioural strategies vary with local breeding synchrony levels remains largely unknown. This is important because, in many species, intra-population patterns of extra-pair fertilisation rates, and hence cuckoldry risk, are known to vary with the number of simultaneously fertile females. Each sex may therefore differ in how they behave towards male conspecifics during different degrees of breeding synchrony. Here I provide evidence of such sex-specific differences in the golden whistler (Pachycephala pectoralis), a species in which within-pair paternity assurance is negatively associated with breeding synchrony. Via simulated territorial intrusions using decoy males, I show that males, but not females, increase levels of aggression to male intruders during periods of low synchrony, possibly because cuckoldry risk is greatest during this period. In addition, males appear to invest more effort into mate guarding after, but not before, territorial intrusions during this period. These inter-sexual differences may reflect conflicts in interest between the sexes, with females consistently showing interest in males during the fertile period regardless of synchrony levels and males investing more resources into expelling intruders when the risk of paternity loss is greatest. This study thus provides evidence that males may be able to detect variation in breeding synchrony and cuckoldry risk and adjust their paternity assurance behaviour accordingly.     

An equal sex ratio followed by differential sex mortality causes overestimation of females in gall midges: no evidence for sex ratio regulation

Monogeny, the production of unisexual broods by individual females, has been recognized for nearly 80?years. The genetic nature of gall midges' sex determination predicts an equal numbers of male-producing and female-producing females in the populations such that the overall sex ratio is expected to be nearly 1:1. However, observations of some strictly monogenous populations with biased sex ratio, mainly toward females, have raised the question of whether gall midges are able to adjust their offspring sex ratio in response to changes in environmental conditions, and some authors have even considered sex ratio regulation as a strong force in the course of the evolution of monogeny. In this paper, first, by studying the sex ratio variations of the predatory gall midge, Aphidoletes aphidimyza within a generation, we showed that adult males emerge up to 1?day earlier and have shorter life span than females (less than 4?days and up to 6?days, respectively). Although, the sex ratio of A. aphidimyza at the time of emergence was nearly 1:1 (52.41?% males), a simple population simulation indicated that the differential mortality of sexes can lead to a female-biased sex ratio estimation (57.88?% females) under random sampling in the natural environments. Our results imply that the primary sex ratio of monogenous gall midges is nearly 1:1 and that the arrhenogenic/thelygenic gall midges are not able to alter the number of their male/female progenies in response to changes in environmental conditions.     

Sex,age, spleen size,and kidney fat of red deer relative to infection intensities of the lungworm <Emphasis Type="Italic">Elaphostrongylus cervi</Emphasis>

We analyzed the relationships among spleen size, body condition (measured as kidney fat), and larval counts of the nematode Elaphostrongylus cervi in red deer (Cervus elaphus). The aim was to investigate the interaction between host body condition and intensity of infection with parasites. As red deer are highly polygynous, we also tested whether these relationships varied with sex and age of the hosts. Kidney fat and spleen size were positively correlated in subadults (2–3 years old) and adults (>3 years old), but not in calves (<1 year old) or yearlings (1–2 years old). Spleen size was negatively associated with nematode load in subadult females and in adult males. These two age classes are potentially the most nutritionally stressed, as subadult hinds are still growing and often engaging in rearing their first calf, and adult stags were sampled just after the rut, which is recognized as a substantial energy drain in this age–sex class, as they compete to hold females during the mating season. Body condition related negatively to parasite count only in adult males. In the context of red deer life history, these findings suggest that spleen size is dependent on body condition and that it could be affected by variation in resource partitioning among immune defense, growth, and reproductive effort in red deer. For the first time in a wild mammal, the spleen mass is shown to be positively related to body condition and negatively related to parasite infection. We conclude that elucidating whether spleen mass reflects immune defense investment or a measure of general body condition should contribute to understanding topical issues in mammal ecology.     

Sperm-depleted males influence the reproductive behaviour of conspecifics

In many insect species, sperm-depleted males (SDMs, i.e. males that have exhausted their sperm after a given number of matings) remain sexually active and continue to mate females. Here, we investigated the behavioural modifications that occur in both sexes of the parasitoid Asobara tabida Nees (Hymenoptera: Braconidae), after matings by fertile males and sperm-depleted males. We show that (i) virgin females, mated females and females mated to a SDM exhibited different behaviours and that (ii) males responded differently to females depending on whether the females had previously mated with an SDM or not. Our findings demonstrate that SDM influenced the reproductive behaviour of both males and females, especially with regard to male responsiveness and female attractiveness. These findings are discussed in the context of adaptive behaviour and fitness maximization in both males and females.     

Geographical variation in parasitism shapes larval immune function in a phytophagous insect

Two of the central goals of immunoecology are to understand natural variation in the immune system among populations and to identify those selection pressures that shape immune traits. Maintenance of the immune system can be costly, and both food quality and parasitism selection pressure are factors potentially driving immunocompetence. In tritrophic interactions involving phytophagous insects, host plants, and natural enemies, the immunocompetence of phytophagous insects is constrained by selective forces from both the host plants and the natural enemies. Here, we assessed the roles of host plants and natural enemies as selective pressures on immune variation among natural populations of Lobesia botrana. Our results showed marked geographical variation in immune defenses and parasitism among different natural populations. Larval immune functions were dependent of the host plant quality and were positively correlated to parasitism, suggesting that parasitoids select for greater investment into immunity in moth. Furthermore, investment in immune defense was negatively correlated with body size, suggesting that it is metabolically expensive. The findings emphasize the roles of host plants and parasitoids as selective forces shaping host immune functions in natural conditions. We argue that kinds of study are central to understanding natural variations in immune functions, and the selective forces beyond.     

Morphometric relationships and reproductive maturation of the mudskipper,Periophthalmus barbarus from subsistence catches in the mangrove swamps of IMO estuary,Nigeria

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