Sex-specific Life History Research Articles

A hypothesis for robust polarization vision: an example from the Australian imperial blue butterfly, Jalmenus evagoras.

The Australian lycaenid butterfly Jalmenus evagoras has iridescent wings that are sexually dimorphic, spectrally and in their degree of polarization, suggesting that these properties are likely to be important in mate recognition. We first describe the results of a field experiment showing that free-flying individuals of J. evagoras discriminate between visual stimuli that vary in polarization content in blue wavelengths but not in others. We then present detailed reflectance spectrophotometry measurements of the polarization content of male and female wings, showing that female wings exhibit blue-shifted reflectance, with a lower degree of polarization relative to male wings. Finally, we describe a novel method for measuring alignment of ommatidial arrays: by measuring variation of depolarized eyeshine intensity from patches of ommatidia as a function of eye rotation, we show that (a) individual rhabdoms contain mutually perpendicular microvilli; (b) many rhabdoms in the array have their microvilli misaligned with respect to neighboring rhabdoms by as much as 45deg; and (c) the misaligned ommatidia are useful for robust polarization detection. By mapping the distribution of the ommatidial misalignments in eye patches of J. evagoras, we show that males and females exhibit differences in the extent to which ommatidia are aligned. Both the number of misaligned ommatidia suitable for robust polarization detection and the number of aligned ommatidia suitable for edge detection vary with respect to both sex and eye patch elevation. Thus, J. evagoras exhibits finely tuned ommatidial arrays suitable for perception of polarized signals, likely to match sex-specific life history differences in the utility of polarized signals.

Philopatry yields higher fitness than dispersal in a cooperative breeder with sex-specific life history trajectories.

Social evolution is tightly linked to dispersal decisions, but the ecological and social factors selecting for philopatry or dispersal often remain obscure. Elucidating selection mechanisms underlying alternative life histories requires measurement of fitness effects in the wild. We report on a long-term field study of 496 individually marked cooperatively breeding fish, showing that philopatry is beneficial as it increases breeding tenure and lifetime reproductive success in both sexes. Dispersers predominantly join established groups and end up in smaller groups when they ascend to dominance. Life history trajectories are sex specific, with males growing faster, dying earlier, and dispersing more, whereas females more likely inherit a breeding position. Increased male dispersal does not seem to reflect an adaptive preference but rather sex-specific differences in intrasexual competition. Cooperative groups may thus be maintained because of inherent benefits of philopatry, of which females seem to get the greater share in social cichlids.

Comparative biology of the grenadiers Macrourus caml and M. whitsoni in the Ross Sea region, Antarctica

The grenadiersMacrourus camlandM. whitsoniform a significant bycatch component of longline fisheries for Antarctic toothfish (Dissostichus mawsoni) in the Southern Ocean. A lack of species-level biological and catch data has to date hindered the development of quantitative assessments for these deepwater species. This paper examines species- and sex-specific life histories ofM.camlandM.whitsoniin the Ross Sea region, Antarctica, from samples collected from the commercial fishery and research collections.Macrourus camlwas found to live longer, grow slower, and attain a larger maximum length thanM.whitsoni, reaching at least 65 years of age and 97 cm total length (TL) vs. 43 years and 78 cm TL forM.whitsoni. In addition,M.camlattains a larger length for a given age and a greater weight for a given length. For each species, females of a given age were larger and reached a greater maximum age than males. Assuming that selectivity of the fishing gear is related to fish size, greater fishing pressure on females than males is likely, and was evidenced by female-biased sex ratios of both species. Estimates of natural and fishing mortality rates were low for both species. Despite having a shorter lifespan and attaining a smaller maximum length,M.whitsonimatured later in life and at larger lengths thanM.caml. Although sampling opportunities are constrained seasonally due to sea ice cover, gonad staging and interannual patterns in gonadosomatic indices suggest prolonged spawning for both species, with available data indicating peak spawning during austral summer. The differences in life history observed between these two closely related and morphologically similar species illustrates the importance of understanding species-specific life histories to infer responses to exploitation, provide key biological inputs to inform parameters for future risk assessments and ecosystem models as well as baseline information for comparative work over both time and space, contribute to greater clarity in managing fisheries that interact with these two species, and further our understanding of grenadier life history.

Sex-Specific Life History Affected by Stocking in Juvenile Brown Trout

Salmonids are a socioeconomically and ecologically important group of fish that are often managed by stocking. Little is known about potential sex-specific effects of stocking, but recent studies found that the sexes differ in their stress tolerances already at late embryonic stage, i.e., before hatchery-born larvae are released into the wild and long before morphological gonad formation. It has also been speculated that sex-specific life histories can affect juvenile growth and mortality, and that a resulting sex-biassed demography can reduce population growth. Here we test whether juvenile brown trout (Salmo trutta) show sex-specific life histories and whether such sex effects differ in hatchery- and wild-born fish. We modified a genetic sexing protocol to reduce false assignment rates and used it to study the timing of sex differentiation in a laboratory setting, and in a large-scale field experiment to study growth and mortality of hatchery- and wild-born fish in different environments. We found no sex-specific mortality in any of the environments we studied. However, females started sex differentiation earlier than males, and while growth rates were similar in the laboratory, they differed significantly in the field depending on location and origin of fish. Overall, hatchery-born males grew larger than hatchery-born females while wild-born fish showed the reverse pattern. Whether males or females grew larger was location-specific. We conclude that juvenile brown trout show sex-specific growth that is affected by stocking and by other environmental factors that remain to be identified.

Caste, Sex, and Parasitism Influence Brain Plasticity in a Social Wasp

Brain plasticity is widespread in nature, as it enables adaptive responses to sensory demands associated with novel stimuli, environmental changes and social conditions. Social Hymenoptera are particularly well-suited to study neuroplasticity, because the division of labor amongst females and the different life histories of males and females are associated with specific sensory needs. Here, we take advantage of the social waspPolistes dominulato explore if brain plasticity is influenced by caste and sex, and the exploitation by the strepsipteran parasiteXenos vesparum.Within sexes, male wasps had proportionally larger optic lobes, while females had larger antennal lobes, which is consistent with the sensory needs of sex-specific life histories. Within castes, reproductive females had larger mushroom body calyces, as predicted by their sensory needs for extensive within-colony interactions and winter aggregations, than workers who frequently forage for nest material and prey. Parasites had different effects on female and male hosts. Contrary to our predictions, female workers were castrated and behaviorally manipulated by female or male parasites, but only showed moderate differences in brain tissue allocation compared to non-parasitized workers. Parasitized males maintained their reproductive apparatus and sexual behavior. However, they had smaller brains and larger sensory brain regions than non-parasitized males. Our findings confirm that caste and sex mediate brain plasticity inP. dominula, and that parasitic manipulation drives differential allocation of brain regions depending on host sex.

Life history, mating dynamics and the origin of parental care.

Parental care, mating dynamics and life history co-evolve. Understanding the diversity of reproductive patterns found in nature is a major focus of evolutionary ecology research. Previous research suggests that the origin of parental care of eggs will be favoured when egg and adult death rates and juvenile survival are relatively high. However, the previous research that explored the link between care and life history did not account for among-species variation in mating dynamics. As mating dynamics are generally expected to influence care, we explore, theoretically, the life-history conditions (stage-specific rates of maturation and survival) that favour parental care across three mating scenarios: reproductive rate (1) is unaffected by males (assuming that some males are present), (2) increases as male abundance increases or (3) decreases as male abundance increases. Across scenarios, all forms of care were most strongly favoured when egg and adult death rates, juvenile survival and female egg maturation rates were relatively high. When reproductive rate was unaffected by male abundance or increased as male abundance increased, as we might expect in systems in which females are mate-limited, all forms of care were most strongly favoured when male egg maturation rate (i.e. the rate at which male eggs develop, mature and hatch) was moderate or high. When greater male abundance inhibited reproduction, which might occur in systems with intense male-male competition, all forms of care were most strongly favoured when male egg maturation rate was low-to-moderate. These results suggest that life history affects the evolution of parental care, and sex-specific life history can interact with mating dynamics to influence the origin of care.

Sex-Specific Stress Tolerance in Embryos of Lake Char (Salvelinus umbla)

Salmonid fish have become important models in evolution and ecology, but possible effects of embryo or larval sex are mostly ignored, probably because morphological gonad formation starts only months after hatching and sexual maturation years later. However, recent gene expression studies and first observations in domestic strains suggest that sex-specific life histories could already start at an embryonic stage. Here we test this hypothesis in embryos and larvae of lake char (Salvelinus umbla). We sampled wild char and used their gametes to produce embryos of 40 different families. Embryos were raised singly in a stress or a non-stress environment until a late larval stage (stress was induced by allowing remainders of ovarian fluids to support microbial growth). Genetic markers were then used to sex the fish and reconstruct paternity (N= 1,463, including dead embryos). Primary sex ratio did not differ among families and was about 1:1. Female embryos hatched on average later and showed lower stress tolerance than male embryos. There were significant parental effects on offspring growth and mortality, but the sex differences in embryo performance were not family specific. We conclude that the sexes differ in their life history and susceptibilities to environmental stress already at embryonic stages. Environmental stress during incubation can therefore affect population sex ratio and hence population growth and genetics.

Changes in life history and population size can explain the relative neutral diversity levels on X and autosomes in extant human populations

In human populations, the relative levels of neutral diversity on the X and autosomes differ markedly from each other and from the naïve theoretical expectation of 3/4. Here we propose an explanation for these differences based on new theory about the effects of sex-specific life history and given pedigree-based estimates of the dependence of human mutation rates on sex and age. We demonstrate that life history effects, particularly longer generation times in males than in females, are expected to have had multiple effects on human X-to-autosome (X:A) diversity ratios, as a result of male-biased mutation rates, the equilibrium X:A ratio of effective population sizes, and the differential responses to changes in population size. We also show that the standard approach of using divergence between species to correct for male mutation bias results in biased estimates of X:A effective population size ratios. We obtain alternative estimates using pedigree-based estimates of the male mutation bias, which reveal that X:A ratios of effective population sizes are considerably greater than previously appreciated. Finally, we find that the joint effects of historical changes in life history and population size can explain the observed X:A diversity ratios in extant human populations. Our results suggest that ancestral human populations were highly polygynous, that non-African populations experienced a substantial reduction in polygyny and/or increase in the male-to-female ratio of generation times around the Out-of-Africa bottleneck, and that current diversity levels were affected by fairly recent changes in sex-specific life history.

Life History Effects on Neutral Diversity Levels of Autosomes and Sex Chromosomes.

Understanding the determinants of neutral diversity patterns on autosomes and sex chromosomes provides a bedrock for the interpretation of population genetic data; in particular, differences between the two informs our understanding of sex-specific demographic and mutation processes. While sex-specific age-structure and variation in reproductive success have long been known to affect neutral diversity, theoretical descriptions of these effects were complicated and lacking in generality, stymying attempts to relate diversity patterns of species with their life history. Here, we derive general yet simple expressions for these effects. In particular, we show that life history effects on X-to-autosome ratios of pairwise diversity levels (X:A diversity ratios) depend only on the male-to-female ratios of mutation rates, generation times, and reproductive variances. Our results reveal that changing the male-to-female ratio of generation times has opposite effects on X:A ratios of diversity and divergence. They also explain how sex-specific life histories modulate the response of X:A diversity ratios to changes in population size. More generally, they clarify that sex-specific life history-generation times in particular-should have marked effects on X:A diversity ratios in many taxa and enable further investigation of these effects.

Sex differences in the impact of social status on hair cortisol concentrations in rhesus monkeys (Macaca mulatta).

Social status impacts stress in primates, but the direction of the effect differs depending on species, social style, and group stability. This complicates our ability to identify broadly applicable principles for understanding how social status impacts health and fitness. One reason for this is the fact that social status is often measured as linear dominance rank, yet social status is more complex than simply high or low rank. Additionally, most research on social status and health ignores the effects of sex and sex-specific relationships, despite known differences in disease risk, coping strategies, and opposite-sex dominance interactions between males and females in many species. We examine the influence of social status, sex, and opposite-sex interactions on hair cortisol concentrations (HCC) in a well-studied species, rhesus macaques, where the literature predicts low ranking individuals would experience more chronic stress. Animals in three captive, seminaturalistic social groups (N = 252; 71 male) were observed for 6 weeks to obtain metrics of social status (rank and dominance certainty [DC]). DC is a measure of one's fit within the hierarchy. Hair samples were collected from each subject and analyzed for HCC. Generalized linear mixed models were used to examine (a) whether rank, DC, or sex predicted HCC; (b) whether same- or opposite-sex dominance relationships differentially impacted HCC; and (c) whether aggressive interactions initiated or received could explain any observed relationships. Results indicated that DC, not rank, predicted HCC in a sex-specific manner. For males, high HCC were predicted by receiving aggression from or having high DC with other males as well as having low DC with females. For females, only high DC with males predicted high HCC. These results likely relate to sex-specific life history pattern differences in inherited versus earned rank that are tied to female philopatry and male immigration.

Signatures of replication timing, recombination, and sex in the spectrum of rare variants on the human X chromosome and autosomes

The sources of human germline mutations are poorly understood. Part of the difficulty is that mutations occur very rarely, and so direct pedigree-based approaches remain limited in the numbers that they can examine. To address this problem, we consider the spectrum of low-frequency variants in a dataset (Genome Aggregation Database, gnomAD) of 13,860 human X chromosomes and autosomes. X-autosome differences are reflective of germline sex differences and have been used extensively to learn about male versus female mutational processes; what is less appreciated is that they also reflect chromosome-level biochemical features that differ between the X and autosomes. We tease these components apart by comparing the mutation spectrum in multiple genomic compartments on the autosomes and between the X and autosomes. In so doing, we are able to ascribe specific mutation patterns to replication timing and recombination and to identify differences in the types of mutations that accrue in males and females. In particular, we identify C > G as a mutagenic signature of male meiotic double-strand breaks on the X, which may result from late repair. Our results show how biochemical processes of damage and repair in the germline interact with sex-specific life history traits to shape mutation patterns on both the X chromosome and autosomes.

Selection for small body size favours contrasting sex-specific life histories, boldness and feeding in medaka, Oryzias latipes

BackgroundStudying variation in life-history traits and correlated behaviours, such as boldness and foraging (i.e., pace-of-life syndrome), allows us to better understand how these traits evolve in a changing environment. In fish, it is particularly relevant studying the interplay of resource abundance and size-selection. These are two environmental stressors affecting fish in natural conditions, but also associated with human-induced environmental change. For instance, fishing, one of the most important threats for freshwater and marine populations, results in both higher mortality on large-sized fish and reduced population density.ResultsMedaka, Oryzias latipes, from lines selected for large or small size over ten generations, were exposed individually to high or low food availability from birth to adulthood. Maturation schedules, reproductive investment, growth, boldness and feeding were assessed to evaluate the effect of size-selection on the pace of life, and whether it differed between food contexts (high and low). Different food abundance and size-selection resulted in diverse life histories associated with different feeding and boldness behaviour, thus showing different pace-of-life-syndromes. High availability of food favoured faster growth, earlier maturation and increased shyness. Selection for small size led to slower growth in both males and females. But, the life-history trajectory to reach such growth was sex- and food-specific. Under low food conditions, females selected for small size showed earlier maturation, which led to slower adult growth and subsequent low willingness to feed, compared to females selected for large size. No line differences were found in females at high food conditions. In contrast, males exposed to selection for small size grew slower both as juvenile and adult, and were bolder under both feeding regimes. Therefore, the response to size-selection was more sensitive to food availability in females than in males.ConclusionsWe showed that size-selection (over ten generations) and resource abundance (over developmental time) led to changes in life history and behaviour. However, the effect of size-selection was sex- and context-specific, calling for precaution when drawing general conclusions on the population-level effects (or lack of them) of size-selective fishing. Conservation and management plans should consider this sex- and context-specificity.

Lifetime Fitness, Sex-Specific Life History, and the Maintenance of a Polyphenism.

Polyphenisms-alternative morphs produced through plasticity-can reveal the evolutionary and ecological processes that initiate and maintain diversity within populations. We examined lifetime fitness consequences of two morphs in a polyphenic population of Arizona tiger salamanders using a 27-year data set with 1,317 adults and 6,862 captures across eight generations. Larval salamanders develop into either an aquatic paedomorph that retains larval traits and stays in its natal pond or a terrestrial metamorph that undergoes metamorphosis. To evaluate the adaptive significance of this polyphenism, we compared lifetime reproductive success of each morph and assessed how life-history strategies and spatiotemporal variation explained fitness. We found sex-specific differences in lifetime fitness between morphs. For males, paedomorphs had more reproductive opportunities than metamorphs when we accounted for the potential mating advantage of larger males. For females, in contrast, metamorphs had higher estimated egg production than paedomorphs. Life-history strategies differed between morphs largely because the morphs maximized different ends of the trade-off between age at first reproduction and longevity. Spatiotemporal variation affected larval more than adult life-history traits, with little to no effect on lifetime fitness. Thus, environmental variation likely explains differences in morph production across time and space but contributes little to lifetime fitness differences between morphs and sexes. Our long-term study and measures of lifetime fitness provide unique insight into the complex selective regimes potentially acting on each morph and sex. Our findings motivate future work to examine how sex-specific selection may contribute to the maintenance of polyphenism.

Rapid evolution of female-biased genes among four species of Anopheles malaria mosquitoes.

Understanding how phenotypic differences between males and females arise from the sex-biased expression of nearly identical genomes can reveal important insights into the biology and evolution of a species. Among Anopheles mosquito species, these phenotypic differences include vectorial capacity, as it is only females that blood feed and thus transmit human malaria. Here, we use RNA-seq data from multiple tissues of four vector species spanning the Anopheles phylogeny to explore the genomic and evolutionary properties of sex-biased genes. We find that, in these mosquitoes, in contrast to what has been found in many other organisms, female-biased genes are more rapidly evolving in sequence, expression, and genic turnover than male-biased genes. Our results suggest that this atypical pattern may be due to the combination of sex-specific life history challenges encountered by females, such as blood feeding. Furthermore, female propensity to mate only once in nature in male swarms likely diminishes sexual selection of post-reproductive traits related to sperm competition among males. We also develop a comparative framework to systematically explore tissue- and sex-specific splicing to document its conservation throughout the genus and identify a set of candidate genes for future functional analyses of sex-specific isoform usage. Finally, our data reveal that the deficit of male-biased genes on the X Chromosomes in Anopheles is a conserved feature in this genus and can be directly attributed to chromosome-wide transcriptional regulation that de-masculinizes the X in male reproductive tissues.

Sex- and cohort-specific life-history strategies in Mongolian gerbils (Meriones unguiculatus)

Information on life-history traits is crucial for understanding rodent evolution, ecology, and behavioral strategies. Here, we report on the results from a 4-year study that collected data on the sexual and seasonal cohort (winter C1, spring C2, and summer C3) life-history traits of Mongolian gerbils (Meriones unguiculatus) maintained in a natural enclosure, using monthly live trapping, in Inner Mongolia (China). We found no sexual differences in the observed longevity, age of maturity, potential reproductive life span, or dispersal in the year of their birth, apart from a male-biased dispersal after gerbils overwintered as adults. Nevertheless, we did find cohort-specific life-history patterns in the above traits; for example, the C1 females matured at a significantly earlier age than did the C2 and C3 females, while the C3 gerbils commonly delayed their reproduction to the following year. Moreover, 21.7% of C1 and 6.1% of C2 females bred more than one litter, yet none of the C3 females did so during their birth-year breeding season. The average number of litters per female of C1 was greater than that of either C2 or C3 in their birth-year breeding season. Our results suggest that sex-specific life history traits may align with the predictions of sexual selection and the monogamy system hypothesis in Mongolian gerbils. In addition, the seasonal cohort-specific life-history traits revealed here lend support to the hypothesis of an adaptive switching behavior—from fast to slow development in the later-born gerbils—as opposed to the hypothesis of direct environmental limitation, although the slight inter-annual variation detected is consistent with the precipitation constraints.

Sex-specific graphs: Relating group-specific topology to demographic and landscape data.

Sex-specific genetic structure is a commonly observed pattern among vertebrate species. Facing differential selective pressures, individuals may adopt sex-specific life history traits that ultimately shape genetic variation among populations. Although differential dispersal dynamics are commonly detected in the literature, few studies have used genetic structure to investigate sex-specific functional connectivity. The recent use of graph theoretic approaches in landscape genetics has demonstrated network capacities to describe complex system behaviours where network topology represents genetic interaction among subunits. Here, we partition the overall genetic structure into sex-specific graphs, revealing different male and female dispersal dynamics of a fisher (Pekania [Martes] pennanti) metapopulation in southern Ontario. Our analyses based on network topologies supported the hypothesis of male-biased dispersal. Furthermore, we demonstrated that the effect of the landscape, identified at the population level, could be partitioned among sex-specific strata. We found that female connectivity was negatively correlated with snow depth, whereas connectivity among males was not. Our findings underscore the potential of conducting sex-specific analysis by identifying landscape elements or configuration that differentially promotes or impedes functional connectivity between sexes, revealing processes that may otherwise remain cryptic. We propose that the sex-specific graph approach would be applicable to other vagile species where differential sex-specific processes are expected to occur.

Differential investment in body girths by sex: Evidence from 3D photonic scanning in a Thai cohort.

Life history trade-offs may manifest between competing organs and tissues in the body. Sexual dimorphism in tissue investment is well-established in humans, with sex-associated body shape differences linked to natural and sexual selection. This study uses three-dimensional (3D) photonic scanning to test whether males and females differentially invest energy in various body regions in relation to two independent proxies of growth. Body shape data (multiple girths) came from a Thai cohort (n = 11,610; 53% female; age range 21-88 years). Weight was considered a proxy for recent energy acquisition. Stature represented completed growth, a proxy for energy acquisition earlier in life. The data were analyzed using growth-proxy by sex interaction log-log regression models adjusting for age, salary and number of children. For a given percentage increase in weight, females showed greater percentage increases than males in girths of the arm, chest, hip, thigh, knee and calf (p < 0.001), whilst males exceeded females in head and waist girths (also p < 0.001). For a given percentage increase in height, weight and all girths showed greater proportional changes in males than females (p < 0.001). These results indicate sex-specific life history strategies wherein the direction and timing of energy investment in girths varies between the sexes. The results add to literature suggesting that sexual dimorphism in body morphology is not a fixed trait; rather, differential energy allocation to specific body regions appears to be a plastic strategy adjusted in relation to energy acquisition across the life course.

The unexpected mating system of the androdioecious barnacle Chelonibia testudinaria (Linnaeus 1758).

Androdioecy was first described by Darwin in his seminal work on barnacle diversity; he identified males and hermaphrodites in the same reproductive population. Today, we realize that many androdioecious plants and animals share astonishing similarities, particularly with regard to their evolutionary history and mating system. Notably, these species were ancestrally dioecious, and their mating system has the following characteristics: hermaphrodites self-fertilize frequently, males are more successful in large mating groups, and males have a mating advantage. A male mating advantage makes androdioecy more likely to persist over evolutionary times. Androdioecious barnacles, however, appear to persist as an outlier with a different evolutionary trajectory: they originate from hermaphroditic species. Although sexual systems of androdioecious barnacles are known, no information on the mating system of androdioecious barnacles is available. This study assessed the mating system of the androdioecious barnacle Chelonibia testudinaria. In contrast to other androdioecious species, C.testudinaria does not self-fertilize, males do not have a mating advantage over hermaphrodites, and the average mating group is quite small, averaging only three individuals. Mating success is increased by proximity to the mate and penis length. Taken together, the mating system of C.testudinaria is unusual in comparison with other androdioecious plants and animals, and the lack of a male mating advantage suggests that the mating system alone does not provide an explanation for the maintenance of androdioecy in this species. Instead, we propose that sex-specific life history equalizes male and hermaphroditic overall fitness.

Life history effects on the molecular clock of autosomes and sex chromosomes

One of the foundational results in molecular evolution is that the rate at which neutral substitutions accumulate on a lineage equals the rate at which mutations arise. Traits that affect rates of mutation therefore also affect the phylogenetic "molecular clock." We consider the effects of sex-specific generation times and mutation rates in species with two sexes. In particular, we focus on the effects that the age of onset of male puberty and rates of spermatogenesis have likely had in hominids (great apes), considering a model that approximates features of the mutational process in mammals, birds, and some other vertebrates. As we show, this model can account for a number of seemingly disparate observations: notably, the puzzlingly low X-to-autosome ratios of substitution rates in humans and chimpanzees and differences in rates of autosomal substitutions among hominine lineages (i.e., humans, chimpanzees, and gorillas). The model further suggests how to translate pedigree-based estimates of human mutation rates into split times among extant hominoids (apes), given sex-specific life histories. In so doing, it largely bridges the gap reported between estimates of split times based on fossil and molecular evidence, in particular suggesting that the human-chimpanzee split may have occurred as recently as 6.6 Mya. The model also implies that the "generation time effect" should be stronger in short-lived species, explaining why the generation time has a major influence on yearly substitution rates in mammals but only a subtle one in human pedigrees.

Dose-dependent effects of an immune challenge at both ultimate and proximate levels in Drosophila melanogaster.

Immune responses are highly dynamic. The magnitude and efficiency of an immune response to a pathogen can change markedly across individuals, and such changes may be influenced by variance in a range of intrinsic (e.g. age, genotype, sex) and external (e.g. abiotic stress, pathogen identity, strain) factors. Life history theory predicts that up-regulation of the immune system will come at a physiological cost, and studies have confirmed that increased investment in immunity can reduce reproductive output and survival. Furthermore, males and females often have divergent reproductive strategies, and this might drive the evolution of sex-specific life history trade-offs involving immunity, and sexual dimorphism in immune responses per se. Here, we employ an experiment design to elucidate dose-dependent and sex-specific responses to exposure to a nonpathogenic immune elicitor at two scales--the 'ultimate' life history and the underlying 'proximate' immune level in Drosophila melanogaster. We found dose-dependent effects of immune challenges on both male and female components of reproductive success, but not on survival, as well as a response in antimicrobial activity. These results indicate that even in the absence of the direct pathogenic effects that are associated with actual disease, individual life histories respond to a perceived immune challenge--but with the magnitude of this response being contingent on the initial dose of exposure. Furthermore, the results indicate that immune responses at the ultimate life history level may indeed reflect underlying processes that occur at the proximate level.