The environment organisms experience during development can have effects which carry over into their adult lives. These environments not only affect adult traits at a given point in time, but also how these traits change with age. Generally, favorable developmental environments lead to more optimal adult traits while stressful environments are deleterious ("silver-spoon effect"). But whether developmental environments affect how whole-organism performance traits change with age or whether they affect males and females differently lacks evidence. Here, we test whether experiencing favorable developmental environments leads to "silver-spoon effects" on life-history traits and whole-organism flight performance of males and females by manipulating developmental density in Callosobruchus maculatus. We further test whether such effects are mediated by an individual's reproductive status by comparing traits of mated versus virgin females. We show that although developmental crowding has no effect on flight performance of either sex, on male adult lifespan, or on female age-dependent survival, it is detrimental to female lifespan and fecundity, as well as female age-dependent reproduction, suggesting that females are more sensitive to differences in developmental density. Additionally, we show that flight traits change with age and differ not only between males and females, which is reflective of sex-specific lifehistories and behavior, but also between virgin and mated females. Our study is important for understanding sex-specific responses to developmental environments in life-history, behavioral, and ecologically important traits which may affect resource allocation, dispersal, and mating strategies, hence fitness.

Reproductive ageing can occur due to the deterioration of both the soma and germline. In males, it has mostly been studied with respect to age-related changes in sperm. However, the somatic component of the ejaculate, seminal fluid, is also essential for maintaining reproductive function. Whilst we know that seminal fluid proteins (SFPs) are required for male reproductive success across diverse taxa, age-related changes in SFP quantity and composition are little understood. Additionally, only few studies have explored the reproductive ageing of the tissues that produce SFPs, and the resulting reproductive outcomes. Here we provide a systematic review of studies addressing how advancing male age affects the production and properties of seminal fluid, in particular SFPs and oxidative stress, highlighting many open questions and generating new hypotheses for further research. We additionally discuss how declines in function of different components of seminal fluid, such as SFPs and antioxidants, could contribute to age-related loss of reproductive ability. Overall, we find evidence that ageing results in increased oxidative stress in seminal fluid and a decrease in the abundance of various SFPs. These results suggest that seminal fluid contributes towards important age-related changes influencing male reproduction. Thus, it is essential to study this mostly ignored component of the ejaculate to understand male reproductive ageing, and its consequences for sexual selection and paternal age effects on offspring.

Developmental and adult environments can interact in complex ways to influence the fitness of individuals. Most studies investigating effects of the environment on fitness focus on environments experienced and traits expressed at a single point in an organism's life. However, environments vary with time, so the effects of the environments that organisms experience at different ages may interact to affect how traits change throughout life. Here, we test whether thermal stress experienced during development leads individuals to cope better with thermal stress as adults. We manipulated temperature during both development and adulthood and measured a range of life-history traits, including senescence, in male and female seed beetles (Callosobruchus maculatus). We found that thermal stress during development reduced adult reproductive performance of females. In contrast, life span and age-dependent mortality were affected more by adult than developmental environments, with high adult temperatures decreasing longevity and increasing age-dependent mortality. Aside from an interaction between developmental and adult environments to affect age-dependent changes in male weight, we did not find any evidence of a beneficial acclimation response to developmental thermal stress. Overall, our results show that effects of developmental and adult environments can be both sex and trait specific, and that a full understanding of how environments interact to affect fitness and ageing requires the integrated study of conditions experienced during different stages of ontogeny.

Senescence, the deterioration of organismal function with advancing age, is a puzzling biological phenomenon. While actuarial senescence (i.e., age-dependent increases in mortality rates) is well described across some taxa, reproductive senescence (i.e.age- dependent declines in reproduction) is less understood, especially in males, with mixed patterns reported across studies. To examine the evidence for male reproductive senescence, we investigated how advancing male age affects ejaculate traits across non-human animals via a meta-analysis yielding 1814 effect sizes from 379 studies. We found no evidence for a general pattern of reproductive senescence. Instead, we found high heterogeneity for how reproduction changes with male age across animals. Some of this heterogeneity (>10%) was associated with biological factors. For example, there were taxonomical differences for some ejaculate traits — sperm motility declined with male age in lab rodents and fish, whereas ejaculate size improved with male age in bulls, fish, and insects. Some methodological factors were also important in explaining this heterogeneity: studies sampling a larger proportion of a species’ lifespan were more likely to detect senescence in ejaculate traits, emphasising the need to examine the full life cycle of species to document senescence. Contrary to predictions, we reveal that the evidence for senescence in ejaculate traits is sporadic. Our findings will help generate novel hypotheses and identify more effective methodological approaches for studying male reproductive senescence.

The environment organisms experience during development can have effects which carry over into their adult lives. These developmental environments may not only affect adult traits at a given point in time, but also how these traits change with age. Generally, stressful developmental environments can lead to sub-optimal adult fitness traits and a faster deterioration of these traits with age. But whether these environments affect how performance traits change with age or whether they affect males and females differently lacks evidence. Here, we test the sex-specific effects of crowding during development on life history traits such as lifespan, fecundity, age-dependent survival, and reproductive senescence, as well as flight performance traits, in the seed beetle- Callosobruchus maculatus. We show that although developmental crowding has no effect on flight performance of either sex, on male adult lifespan, or on female age-dependant survival, it is detrimental to female lifespan and fecundity, as well as female age-dependent reproduction. Additionally, we show that flight traits can change with age and differ between males and females, which may be reflective of sex-specific life-histories and behaviour. Our study has implications for understanding how developmental environments affect life history and behavioural strategies of both males and females.