Striatal dopamine, sexual activity and lifespan. Longevity of rats treated with (-)deprenyl

The influence of longterm deprenyl treatment on the sexual performance and lifespan of male rats was studied. One hundred and thirty two rats were treated from the end of their 2nd year of life either with saline (1 ml/kg, s.c.) (n = 66) or with deprenyl (0.25 mg/kg, s.c.) (n = 66) three times a week until death. Whereas none of the two-year-old saline-treated rats displayed full scale sexual activity, this appeared in 64 out of 66 rats on deprenyl. The longest living rat in the saline-treated group lived 164 weeks. The lifespan of the group was 147.05 +/- 0.56 weeks. The shortest living animal in the (-)deprenyl-treated group lived 171 weeks and the longest living rat died during the 226th week of its life. The lifespan was 191.91 +/- 2.31 weeks. This is the first instance that a well aimed medication prolonged lifespan of members of a species beyond their maximum age of death (182 weeks in the rat). A close relation between sexual activity and lifespan was detected.     

The Evolution of Senescence and Post-Reproductive Lifespan in Guppies (Poecilia reticulata)

The study of post-reproductive lifespan has been of interest primarily with regard to the extended post-menopausal lifespan seen in humans. This unusual feature of human demography has been hypothesized to have evolved because of the “grandmother” effect, or the contributions that post-reproductive females make to the fitness of their children and grandchildren. While some correlative analyses of human populations support this hypothesis, few formal, experimental studies have addressed the evolution of post-reproductive lifespan. As part of an ongoing study of life history evolution in guppies, we compared lifespans of individual guppies derived from populations that differ in their extrinsic mortality rates. Some of these populations co-occur with predators that increase mortality rate, whereas other nearby populations above barrier waterfalls are relatively free from predation. Theory predicts that such differences in extrinsic mortality will select for differences in the age at maturity, allocation of resources to reproduction, and patterns of senescence, including reproductive declines. As part of our evaluation of these predictions, we quantified differences among populations in post-reproductive lifespan. We present here the first formal, comparative study of the evolution of post-reproductive lifespan as a component of the evolution of the entire life history. Guppies that evolved with predators and that experienced high extrinsic mortality mature at an earlier age but also have longer lifespans. We divided the lifespan into three non-overlapping components: birth to age at first reproduction, age at first reproduction to age at last reproduction (reproductive lifespan), and age at last reproduction to age at death (post-reproductive lifespan). Guppies from high-predation environments live longer because they have a longer reproductive lifespan, which is the component of the life history that can make a direct contribution to individual fitness. We found no differences among populations in post-reproductive lifespan, which is as predicted since there can be no contribution of this segment of the life history to an individual's fitness. Prior work on the evolution of post-reproductive lifespan has been dominated by speculation and correlative analyses. We show here that this component of the life history is accessible to formal study as part of experiments that quantify the different segments of an individual's life history. Populations of guppies subject to different mortality pressures from predation evolved differences in total lifespan, but not in post-reproductive lifespan. Rather than showing the direct effects of selection characterizing other life-history traits, post-reproductive lifespan in these fish appears to be a random add-on at the end of the life history. These findings support the hypothesis that differences in lifespan evolving in response to selection are confined to the reproductive lifespan, or those segments of the life history that make a direct contribution to fitness. We also show, for the first time, that fish can have reproductive senescence and extended post-reproductive lifespans despite the general observation that they are capable of producing new primary oocytes throughout their lives.     

Life expectancy,maximum longevity and lifetime reproductive success in female Thornicroft's giraffe in Zambia

Gestation and longevity scale with body mass across taxa, yet within size dimorphic taxa, males tend to have reduced lifespans compared with females. Testing life history models, and accounting for sex differences in longevity, requires obtaining accurate longitudinal data from wild populations. We provide the first report describing key life history parameters from a long‐term study of giraffes in Africa. We followed a population of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) in Zambia for over 40 years. Maximum longevity among females was approximately 28 years, with lifespan accounting for 81% of the variance in lifetime reproductive success. Average adult female life expectancy was no different than average adult male life expectancy. However, the breeding lifespan of males was about half that of females, while maximum lifespan of males was 75% that of females. Our findings support the suggestion that sex differences in maximum lifespan arise from stronger selection for lengthy lives in females than in males. Among females, longer lives are associated with greater reproductive output.     

The evolution of senescence and post-reproductive lifespan in guppies (Poecilia reticulata)

The study of post-reproductive lifespan has been of interest primarily with regard to the extended post-menopausal lifespan seen in humans. This unusual feature of human demography has been hypothesized to have evolved because of the “grandmother” effect, or the contributions that post-reproductive females make to the fitness of their children and grandchildren. While some correlative analyses of human populations support this hypothesis, few formal, experimental studies have addressed the evolution of post-reproductive lifespan. As part of an ongoing study of life history evolution in guppies, we compared lifespans of individual guppies derived from populations that differ in their extrinsic mortality rates. Some of these populations co-occur with predators that increase mortality rate, whereas other nearby populations above barrier waterfalls are relatively free from predation. Theory predicts that such differences in extrinsic mortality will select for differences in the age at maturity, allocation of resources to reproduction, and patterns of senescence, including reproductive declines. As part of our evaluation of these predictions, we quantified differences among populations in post-reproductive lifespan. We present here the first formal, comparative study of the evolution of post-reproductive lifespan as a component of the evolution of the entire life history.

Guppies that evolved with predators and that experienced high extrinsic mortality mature at an earlier age but also have longer lifespans. We divided the lifespan into three non-overlapping components: birth to age at first reproduction, age at first reproduction to age at last reproduction (reproductive lifespan), and age at last reproduction to age at death (post-reproductive lifespan). Guppies from high-predation environments live longer because they have a longer reproductive lifespan, which is the component of the life history that can make a direct contribution to individual fitness. We found no differences among populations in post-reproductive lifespan, which is as predicted since there can be no contribution of this segment of the life history to an individual's fitness.

Prior work on the evolution of post-reproductive lifespan has been dominated by speculation and correlative analyses. We show here that this component of the life history is accessible to formal study as part of experiments that quantify the different segments of an individual's life history. Populations of guppies subject to different mortality pressures from predation evolved differences in total lifespan, but not in post-reproductive lifespan. Rather than showing the direct effects of selection characterizing other life-history traits, post-reproductive lifespan in these fish appears to be a random add-on at the end of the life history. These findings support the hypothesis that differences in lifespan evolving in response to selection are confined to the reproductive lifespan, or those segments of the life history that make a direct contribution to fitness. We also show, for the first time, that fish can have reproductive senescence and extended post-reproductive lifespans despite the general observation that they are capable of producing new primary oocytes throughout their lives.

     

Fecundity, lifespan and egg mass in butterflies: effects of male-derived nutrients and female size

1. Effects of larval reserves and nutrients received as adults on fecundity and lifespan in female Danaus plexippus (the Monarch Butterfly) were measured to determine the relative importance of different sources of nutrients for reproduction and somatic maintenance.
2. Egg-laying lifespan was correlated with female size but not with the amount of male-derived nutrients or adult food concentration.
3. Lifetime fecundity was higher when females received a large first spermatophore, but was not affected by female size when lifespan was controlled or by adult food concentration.
4. At the end of their lives, females contained unlaid eggs and retained, on average, 88% of their initial mass. This proportion was unchanged in two years, although mean egg-laying lifespan varied from 22·5 to 28·7 days.
5. Egg mass decreased over the female lifespan, and was correlated with female size.
6. These results suggest that larval reserves are more important for somatic maintenance than adult income, but that the protein-rich nutrients received from males contribute to egg production. This supports theoretical predictions and empirical studies of other Lepidoptera showing that larval reserves are less likely to affect fecundity when the adult income can contribute substantially to egg production.     

The evolution of aging and age-related physical decline in mice selectively bred for high voluntary exercise

We tested whether selective breeding for early-age high voluntary exercise behavior over 16 generations caused the evolution of lifelong exercise behavior, life expectancy, and age-specific mortality in house mice (Mus domesticus). Sixteenth-generation mice from four replicate selection lines and four replicate random-bred control lines were individually housed from weaning through death and divided between two activity treatments (either with or without running wheels). Thus, there were four treatment groups: selection versus control crossed with active versus sedentary. The effects of selective breeding on life expectancy and age-specific mortality differed between females and males. In females, sedentary selection mice had early and high initial adult mortality and thus the lowest increases in mortality with age. Active selection females had the lowest early adult mortality, had limited mortality during midlife, and exhibited rapid increases in mortality rates at the very end of life; thus, they had deferred senescence. Median life expectancy was greater for both groups of selection females than for the two complementary groups of control females. Like females, sedentary selection males had the highest early adult mortality, and slow but steadily increasing mortality over the entire lifetime. Unlike the active selection females, active control males had the lowest mortality across the lifespan (until the end of life). Interestingly, the males with the lowest median life expectancy were those in the active selection treatment group. In both sexes, running (km/week) decreased over the lifetime to very low and virtually equivalent levels at the end of life in control and selection mice. Overall, these results demonstrate an evolutionary cost of selective breeding for males, regardless of exercise level, but a benefit for females when they have an outlet for the up-selected behavior. We conclude that correlated evolution of senescence occurs in mice selectively bred for high voluntary wheel running; exercise per se is beneficial for control mice of both sexes, but the impact on the effect of selection depends on sex; and the behavioral effect of exercise selection at an early age declines throughout the life span, which demonstrates decreasing genetic correlations over age for the genes involved in increased exercise.     

Evolution of gut microbial community through reproductive life in female rabbits and investigation of the link with offspring survival

The digestive microbiota plays a decisive role in shaping and preserving health throughout life. Rabbit younglings are born with a sterile digestive tract but then it gets progressively colonised by the microbiota of the nursing mother, by entering in contact with or ingesting the maternal droppings present in the nest. Here we posit that (i) offspring survival and (ii) lifespan of female rabbits are linked to how diverse their microbiota are. To test the hypothesis that maternal microbiota evolves in females having had different levels of offspring survival in their lifetime, we obtained 216 hard faecal samples from 75 female rabbits at ages 19.6, 31.6, 62.6 and 77.6 weeks. The annual mean offspring survival (MOS) at 64 days was calculated for each female then crossed against three alpha-diversity indexes (operational taxonomic units (OTUs), inverse Simpson index and Shannon index). Age was also analysed against these three parameters. The alpha-diversity indexes of the female faecal microbiota did not correlate with MOS, but they did decrease with age (e.g. from 712 OTUs at age 19.6 weeks to 444 OTUs at 77.6 weeks; P < 0.05). The age effect was also found in beta-diversity non-metric multidimensional scaling plots using the Bray–Curtis dissimilarity index and the unweighted UniFrac index but not for MOS. The ability of the microbiota composition from the faecal samples of young females (19.6 weeks old) to predict their lifespan was also evaluated. After subdividing the initial population into two classes (females that weaned a maximum of three litters and females living longer), we found no clear distinction between these two classes. To our knowledge, this is the first long-term study to characterise the gut microbiota of adult female rabbits through their reproductive life, thus laying foundations for using the gut microbiota data and its influence in studies on adult rabbits.     

Life History of Cercopithecus mitis stuhlmanni in the Kakamega Forest, Kenya

Comparative data from wild populations are necessary to understand the evolution of primate life history strategies. We present demographic data from a 29-yr longitudinal study of 8 groups of individually recognized wild blue monkeys (Cercopithecus mitis stuhlmanni). We provide estimates of life history variables and a life table for females. Most females had their first infant at 7 yr. The mean interbirth interval was 28 mo, and decreased from 31 to 18 mo if the first infant died within a year. Interbirth intervals did not differ according to infant sex, but females had longer intervals after their first vs. subsequent births. Infant mortality was 23% and did not differ strongly by sex or mother’s parity. Maximal female lifespan was 32.5–34.5 yr. Across the lifespan, both survivorship and fecundity showed typical primate patterns. Survivorship was lowest in infants, leveled off among juveniles, and then decreased gradually with increasing age in later life. Fecundity was highest among young females and decreased among older females. Births were seasonal, with 64% occurring within 3 mo at the end of the dry season and beginning of the wet season. Survival to 12 mo was higher for infants born during drier months. Birth season timing is plausibly related to thermoregulation of infants, weanling foods, or maternal energy demand. Blue monkeys are a forest-dependent species with a very slow life history and relatively low immature and adult mortality rates compared to closely related guenons living in open habitats. Even among cercopithecines as a whole, they appear to have an exceptionally slow life history relative to body size. Differences in life history “speed” between blue monkeys and their close relatives seem to be related to lower juvenile and adult mortality in forests relative to more open habitats.     

Adaptation to developmental diet influences the response to selection on age at reproduction in the fruit fly

Experimental evolution (EE) is a powerful tool for addressing how environmental factors influence life‐history evolution. While in nature different selection pressures experienced across the lifespan shape life histories, EE studies typically apply selection pressures one at a time. Here, we assess the consequences of adaptation to three different developmental diets in combination with classical selection for early or late reproduction in the fruit fly Drosophila melanogaster. We find that the response to each selection pressure is similar to that observed when they are applied independently, but the overall magnitude of the response depends on the selection regime experienced in the other life stage. For example, adaptation to increased age at reproduction increased lifespan across all diets; however, the extent of the increase was dependent on the dietary selection regime. Similarly, adaptation to a lower calorie developmental diet led to faster development and decreased adult weight, but the magnitude of the response was dependent on the age‐at‐reproduction selection regime. Given that multiple selection pressures are prevalent in nature, our findings suggest that trade‐offs should be considered not only among traits within an organism, but also among adaptive responses to different—sometimes conflicting—selection pressures, including across life stages.     

Longevity determined by developmental arrest genes in Caenorhabditis elegans

The antagonistic pleiotropy theory of aging proposes that aging takes place because natural selection favors genes that confer benefit early on life at the cost of deterioration later in life. This theory predicts that genes that impact development would play a key role in shaping adult lifespan. To better understand the link between development and adult lifespan, we examined the genes previously known to be essential for development. From a pool of 57 genes that cause developmental arrest after inhibition using RNA interference, we have identified 24 genes that extend lifespan in Caenorhabditis elegans when inactivated during adulthood. Many of these genes are involved in regulation of mRNA translation and mitochondrial functions. Genetic epistasis experiments indicate that the mechanisms of lifespan extension by inactivating the identified genes may be different from those of the insulin/insulin-like growth factor 1 (IGF-1) and dietary restriction pathways. Inhibition of many of these genes also results in increased stress resistance and decreased fecundity, suggesting that they may mediate the trade-offs between somatic maintenance and reproduction. We have isolated novel lifespan-extension genes, which may help understand the intrinsic link between organism development and adult lifespan.     

Effects of Increased Flight on the Energetics and Life History of the Butterfly Speyeria mormonia

Movement uses resources that may otherwise be allocated to somatic maintenance or reproduction. How does increased energy expenditure affect resource allocation? Using the butterfly Speyeria mormonia, we tested whether experimentally increased flight affects fecundity, lifespan or flight capacity. We measured body mass (storage), resting metabolic rate and lifespan (repair and maintenance), flight metabolic rate (flight capacity), egg number and composition (reproduction), and food intake across the adult lifespan. The flight treatment did not affect body mass or lifespan. Food intake increased sufficiently to offset the increased energy expenditure. Total egg number did not change, but flown females had higher early-life fecundity and higher egg dry mass than control females. Egg dry mass decreased with age in both treatments. Egg protein, triglyceride or glycogen content did not change with flight or age, but some components tracked egg dry mass. Flight elevated resting metabolic rate, indicating increased maintenance costs. Flight metabolism decreased with age, with a steeper slope for flown females. This may reflect accelerated metabolic senescence from detrimental effects of flight. These effects of a drawdown of nutrients via flight contrast with studies restricting adult nutrient input. There, fecundity was reduced, but flight capacity and lifespan were unchanged. The current study showed that when food resources were abundant, wing-monomorphic butterflies living in a continuous meadow landscape resisted flight-induced stress, exhibiting no evidence of a flight-fecundity or flight-longevity trade-off. Instead, flight changed the dynamics of energy use and reproduction as butterflies adopted a faster lifestyle in early life. High investment in early reproduction may have positive fitness effects in the wild, as long as food is available. Our results help to predict the effect of stressful conditions on the life history of insects living in a changing world.     

A trade-off between female lifespan and larval diet breadth at the interspecific level in Lepidoptera

A prediction arising from several evolutionary diet breadth models is that, in insect herbivores whose adults practise adaptive host plant selection based on larval performance, female adult lifespan should be negatively correlated with larval diet breadth. In one category of models, female adult lifespan drives evolutionary changes in larval diet breadth; in the other category, larval diet breadth drives evolutionary changes in female adult lifespan. Applying the method of independent contrasts to a biologically and phylogenetically diverse array of Lepidoptera, we ask whether larval diet breadth—as measured by the number of larval food plant species reported in the literature—is negatively correlated with female adult lifespan at the interspecific level. We show that these two life history variables are indeed inversely related. Next, we relax the assumption, common to all of the models, that the female adult is the life stage responsible for the distribution of progeny among different host plants. By introducing into our data set three species whose females are incapable of flight (due to either aptery or brachyptery), and whose larvae are the dispersive stage, the negative correlation between female adult lifespan and larval diet breadth is lost, when using the independent contrasts method. We interpret this effect as supporting the models’ common prediction. Ours is the first reported evidence of a lifespan/diet breadth trade-off at the interspecific level among insects, and it confirms the findings of a previous study in which the degree of habitat specialisation among arthropods was inversely related to proxy measures of the degree of search time constraint. In one of our “diet breadth drives changes in lifespan” models, the females’ type of egg maturation strategy (as measured by the ovigeny index) is predicted to be positively correlated with larval diet breadth, and it mediates a female adult lifespan/larval diet breadth trade-off; however, we found no convincing support for such a role.     

Complex genetic architecture of population differences in adult lifespan of a beetle: nonadditive inheritance, gender differences, body size and a large maternal effect

Evolutionary responses to selection can be complicated when there is substantial nonadditivity, which limits our ability to extrapolate from simple models of selection to population differentiation and speciation. Studies of Drosophila melanogaster indicate that lifespan and the rate of senescence are influenced by many genes that have environment- and sex-specific effects. These studies also demonstrate that interactions among alleles (dominance) and loci (epistasis) are common, with the degree of interaction differing between the sexes and among environments. However, little is known about the genetic architecture of lifespan or mortality rates for organisms other than D. melanogaster. We studied genetic architecture of differences in lifespan and shapes of mortality curves between two populations of the seed beetle, Callosobruchus maculatus (South India and Burkina Faso populations). These two populations differ in various traits (such as body size and adult lifespan) that have likely evolved via host-specific selection. We found that the genetic architecture of lifespan differences between populations differs substantially between males and females; there was a large maternal effect on male lifespan (but not on female lifespan), and substantial dominance of long-life alleles in females (but not males). The large maternal effect in males was genetically based (there was no significant cytoplasmic effect) likely due to population differences in maternal effects genes that influence lifespan of progeny. Rearing host did not affect the genetic architecture of lifespan, and there was no evidence that genes on the Y-chromosome influence the population differences in lifespan. Epistatic interactions among loci were detectable for the mortality rate of both males and females, but were detectable for lifespan only after controlling for body size variation among lines. The detection of epistasis, dominance, and sex-specific genetic effects on C. maculatus lifespan is consistent with results from line cross and quantitative trait locus studies of D. melanogaster.     

Adaptive variation in senescence: reproductive lifespan in a wild salmon population

The antagonistic pleiotropy theory of senescence postulates genes or traits that have opposite effects on early-life and late-life performances. Because selection is generally weaker late in life, genes or traits that improve early-life performance but impair late-life performance should come to predominate. Variation in the strength of age-specific selection should then generate adaptive variation in senescence. We demonstrate this mechanism by comparing early and late breeders within a population of semelparous capital-breeding sockeye salmon (Oncorhynchus nerka). We show that early breeders (but not late breeders) are under strong selection for a long reproductive lifespan (RLS), which facilitates defence of their nests against disturbance by later females. Accordingly, early females invest less energy in egg production while reserving more for nest defence. Variation along this reproductive trade-off causes delayed or slower senescence in early females (average RLS of 26 days) than in late females (reproductive lifespan of 12 days). We use microsatellites to confirm that gene flow is sufficiently limited between early and late breeders to allow adaptive divergence in response to selection. Because reproductive trade-offs should be almost universal and selection acting on them should typically vary in time and space, the mechanism described herein may explain much of the natural variation in senescence.     

EFFECTS ON FITNESS COMPONENTS OF P-ELEMENT INSERTS IN DROSOPHILA MELANOGASTER: ANALYSIS OF TRADE-OFFS

We analyzed the trade-offs between fitness components detected in four experiments in which traits were manipulated by inserting small (control) and large (treatment) P-elements into the Drosophila melanogaster genome. Treatment effects and the interactions of treatment with temperature, experiment, and line were caused by the greater length and different positions of the treatment insert. In inbred flies, the treatment decreased early and total fecundity. Whether it increased the lifespan of mated females depended upon adult density. Analysis of line-by-treatment-by-temperature interactions revealed hidden trade-offs that would have been missed by other methods. They included a significant trade-off between lifespan and early fecundity. At 25°C high early fecundity was associated with decreased reproductive rates and increased mortality rates 10–15 days later and persisting throughout life, but not at 29.5°C. Correlations with Gompertz coefficients suggested that flies that were heavier at eclosion also aged more slowly and that flies that aged more slowly had higher fecundity late in life at 25°C. The results support the view that lifespan trades off with fecundity and that late fecundity trades off with rate of aging in fruitflies. Genetic engineering is an independent method for the analysis of trade-offs that complements selection experiments.     

Life history in male mandrills (Mandrillus sphinx): physical development, dominance rank, and group association

We assess life history from birth to death in male mandrills (Mandrillus sphinx) living in a semifree-ranging colony in Gabon, using data collected for 82 males that attained at least the age of puberty, including 33 that reached adulthood and 25 that died, yielding data for their entire lifespan. We describe patterns of mortality and injuries, dominance rank, group association, growth and stature, and secondary sexual character expression across the male lifespan. We examine relationships among these variables and investigate potential influences on male life history, including differences in the social environment (maternal rank and group demography) and early development, with the aim of identifying characteristics of successful males. Sons of higher-ranking females were more likely to survive to adulthood than sons of low-ranking females. Adolescent males varied consistently in the rate at which they developed, and this variation was related to a male's own dominance rank. Males with fewer peers and sons of higher-ranking and heavier mothers also matured faster. However, maternal variables were not significantly related to dominance rank during adolescence, the age at which males attained adult dominance rank, or whether a male became alpha male. Among adult males, behavior and morphological development were related to a male's own dominance rank, and sons of high-ranking females were larger than sons of low-ranking females. Alpha males were always the most social, and the most brightly colored males, but were not necessarily the largest males present. Finally, alpha male tenure was related to group demography, with larger numbers of rival adult males and maturing adolescent males reducing the time a male spent as alpha male. Tenure did not appear to be related to characteristics of the alpha male himself.     

How Does Male Ritual Behavior Vary Across the Lifespan?

Ritual behaviors of some form exist in every society known to anthropologists. Despite this universality, we have little understanding of how ritual behavior varies within populations or across the lifespan, nor the determinants of this variation. Here we test hypotheses derived from life history theory by using behavioral observations and oral interview data concerning participant variation in Fijian kava-drinking ceremonies. We predicted that substantial variation in the frequency and duration of participation would result from (1) trade-offs with reproduction and (2) the intrinsic status differences between ritual participants. We demonstrate that when controlling for household composition, men with young offspring participated less frequently and exhibited greater variance in their time spent at ceremonies than men without young children. However, men with a larger number of total dependents in their household participated more frequently than those with fewer. Moreover, we found that men’s ascribed rank, level of education, and reliance on wage labor all significantly predict their frequency of attendance. We also found that the number of dependents a man has in his household is positively correlated with total food production, and the amount of kava he cultivates. In general, these results suggest that ritual participation is part of an important strategy employed by Fijian men for both achieving status and developing social alliances. Variation in participation in kava ceremonies by Fijian men therefore reflects the constraints of their current life history condition and their inherited rank.     

Pathways to fitness: carry‐over effects of late hatching and urbanisation on lifetime mating success

Life history theory and most empirical studies assume carry‐over effects of larval ­conditions to shape adult fitness through their impact on metamorphic traits (age and mass at metamorphosis). Yet, very few formal tests of this connection across metamorphosis exist, because this entails longitudinal studies from the egg stage and requires measuring fitness in (semi)natural conditions. In a longitudinal one‐year common‐garden rearing experiment consisting of an outdoor microcosm part for the larval stage and a large outdoor insectary part for the adult stage, we studied the effects of two factors related to time constraints in the larval stage (egg hatching period and urbanisation) on life history traits and lifetime mating success in the males of the damselfly Coenagrion puella. We reared early‐ and late‐hatched larvae from each of three rural and three urban populations from the egg stage throughout their adult life. Key findings were that both the hatching period and urbanisation shaped adult fitness, yet through different pathways. As expected, the more time‐constrained late‐hatched individuals accelerated their larval life history and this was associated with a lower lifetime mating success. A path analysis revealed this carry‐over effect was mediated by the changes in the two metamorphic traits (reduced age and lower mass at emergence). Notably, urban males had a 50% lower lifetime mating success, which was not mediated by age and mass at emergence, and possibly driven by their shorter lifespan. Our results point to long‐term carry‐over effects of the usually ignored natural variation in egg hatching dates, and further contribute to the limited evidence showing fitness costs of adjusting to an urban lifestyle.     

Local adaptations of life‐history traits of a Drosophila parasitoid,Leptopilina boulardi: does climate drive evolution?

1. Climate is an important source of selection on life histories, and local adaptations to climate have been described in several cline studies. Temperature is the main climatic factor that has been considered as an agent of selection, whereas other factors may vary with it, such as precipitation. 2. We compared life‐history traits of five populations of Leptopilina boulardi, a Drosophila parasitoid, originating from contrasting climates. Referring to cline studies, we hypothesised shorter lifespan, earlier reproduction, and lower lipid content in populations from the hottest and driest areas if life histories have been selected in response to temperature and/or humidity. 3. Our results are opposite to these predictions. Females from humid and mild climates invested more in early reproduction and lived for fewer days than females from dry and hot areas, which were synovigenic (i.e. they matured additional eggs during adult life) and able to synthesise lipids during adult life. 4. We suggest that life histories are more adapted to host distribution than to climatic factors. Drosophila patches are more abundant in the humid area, allowing the parasitoids to spend less energy and time finding hosts. This may result in selection for early reproduction traded‐off against longevity. In the hot and dry climate, females have to fly large distances to find host patches. Synovigeny, a long lifespan, lipogenesis, and high dispersal ability may be adaptive there. This is the first time that between‐population differences in the ability to synthesise lipids have been described in parasitoids.     

Why men matter: mating patterns drive evolution of human lifespan

Evolutionary theory predicts that senescence, a decline in survival rates with age, is the consequence of stronger selection on alleles that affect fertility or mortality earlier rather than later in life. Hamilton quantified this argument by showing that a rare mutation reducing survival is opposed by a selective force that declines with age over reproductive life. He used a female-only demographic model, predicting that female menopause at age ca. 50 yrs should be followed by a sharp increase in mortality, a "wall of death." Human lives obviously do not display such a wall. Explanations of the evolution of lifespan beyond the age of female menopause have proven difficult to describe as explicit genetic models. Here we argue that the inclusion of males and mating patterns extends Hamilton's theory and predicts the pattern of human senescence. We analyze a general two-sex model to show that selection favors survival for as long as men reproduce. Male fertility can only result from matings with fertile females, and we present a range of data showing that males much older than 50 yrs have substantial realized fertility through matings with younger females, a pattern that was likely typical among early humans. Thus old-age male fertility provides a selective force against autosomal deleterious mutations at ages far past female menopause with no sharp upper age limit, eliminating the wall of death. Our findings illustrate the evolutionary importance of males and mating preferences, and show that one-sex demographic models are insufficient to describe the forces that shape human senescence.