Multiple simultaneous treatments change plant response from adaptive parental effects to within‐generation plasticity,in Arabidopsis thaliana

In general, studies on plant phenotypic plasticity concentrate on plant responses to different levels of a single environmental factor. Under natural conditions, however, multiple environmental factors often vary simultaneously. I studied the consequences for lifetime fitness caused by single treatments or treatment combinations by investigating patterns of phenotypic plasticity within and between generations. The parental plants (three genotypes of the annual plant Arabidopsis thaliana) received zero, one or two stress treatments at an early life‐stage. The treatments included wounding, shading, chilling, and their pairwise combinations. In the second generation, offspring of treated plants received either the parental or no treatment. Offspring of non‐treated plants were reared under all treatment conditions. Plants responded strongly to the treatments, especially through delayed reproduction, which positively affected lifetime fitness. Notably, treatment combinations triggered stronger plastic responses on average. Because the delay in reproduction was offset by a delay in senescence, the treatments resulted in a fitness gain instead of a loss. However, under adverse environmental conditions, this delay represents a potential fitness cost, especially when the time for reproduction is limited. The treatments ‘wounding’ and ‘shading’ triggered parental effects that increased fitness only in plants that themselves received the treatment. Untreated offspring of wounded or shaded parents performed like control plants. Also, these parental effects were not accompanied by potential fitness costs, such as delayed reproduction. Chilling triggered genotype‐specific parental effects that increased or reduced fitness. Of the treatment combinations only ‘wounding’ and ‘shading’ resulted in genotype‐specific parental effects that increased or reduced fitness independently of offspring treatment. These results suggest that the response of annual plants to treatment combinations triggers predominantly within‐generation plastic responses that include potential fitness costs, which cannot be inferred from studies that manipulate environmental factors individually. Therefore, single treatment studies likely underestimate the costs of plasticity in natural environments.     

Sex‐specific pathways of parental age effects on offspring lifetime reproductive success in a long‐lived seabird

The conditions under which individuals are reared vary and sensitivity of offspring to such variation is often sex‐dependent. Parental age is one important natal condition with consequences for aspects of offspring fitness, but reports are mostly limited to short‐term fitness consequences and do not take into account offspring sex. Here we used individual‐based data from a large colony of a long‐lived seabird, the common tern Sterna hirundo, to investigate longitudinal long‐term fitness consequences of parental age in relation to both offspring and parental sex. We found that recruited daughters from older mothers suffered from reduced annual reproductive success. Recruited sons from older fathers were found to suffer from reduced life span. Both effects translated to reductions in offspring lifetime reproductive success. Besides revealing novel sex‐specific pathways of transgenerational parental age effects on offspring fitness, which inspire studies of potential underlying mechanisms, our analyses show that reproductive senescence is only observed in the common tern when including transgenerational age effects. In general, our study shows that estimates of selective pressures underlying the evolution of senescence, as well as processes such as age‐dependent mate choice and sex allocation, will depend on whether causal transgenerational effects exist and are taken into account.     

Offspring of older parents are smaller—but no less bilaterally symmetrical—than offspring of younger parents in the aquatic plant Lemna turionifera

Offspring quality decreases with parental age in many taxa, with offspring of older parents exhibiting reduced life span, reproductive capacity, and fitness, compared to offspring of younger parents. These “parental age effects,” whose consequences arise in the next generation, can be considered as manifestations of parental senescence, in addition to the more familiar age‐related declines in parent‐generation survival and reproduction. Parental age effects are important because they may have feedback effects on the evolution of demographic trajectories and longevity. In addition to altering the timing of offspring life‐history milestones, parental age effects can also have a negative impact on offspring size, with offspring of older parents being smaller than offspring of younger parents. Here, we consider the effects of advancing parental age on a different aspect of offspring morphology, body symmetry. In this study, we followed all 403 offspring of 30 parents of a bilaterally symmetrical, clonally reproducing aquatic plant species, Lemna turionifera, to test the hypothesis that successive offspring become less symmetrical as their parent ages, using the “Continuous Symmetry Measure” as an index. Although successive offspring of aging parents older than one week became smaller and smaller, we found scant evidence for any reduction in bilateral symmetry.     

PARENTAL AGE,GAMETIC AGE,AND INBREEDING INTERACT TO MODULATE OFFSPRING VIABILITY IN DROSOPHILA MELANOGASTER

In principle, parental relatedness, parental age, and the age of parental gametes can all influence offspring fitness through inbreeding depression and the parental effects of organismal and postmeiotic gametic senescence. However, little is known about the extent to which these factors interact and contribute to fitness variation. Here, we show that, in Drosophila melanogaster, offspring viability is strongly affected by a three‐way interaction between parental relatedness, parental age, and gametic age at successive developmental stages. Overall egg‐to‐adult viability was lowest for offspring produced with old gametes of related, young parents. This overall effect was largely determined at the pupa–adult stage, although three‐way interactions between parental relatedness, parental age and gametic age also explained variation in egg hatchability and larva‐pupa survival. Controlling for the influence of parental and gametic age, we show that inbreeding depression is negligible for egg hatchability but significant at the larva–pupa and pupa–adult stages. At the pupa–adult stage, where offspring could be sexed, parental relatedness, parental age, and gametic age interacted differently in male and female offspring, with daughters suffering higher inbreeding depression than sons. Collectively, our results demonstrate that the architecture of offspring fitness is strongly influenced by a complex interaction between parental effects, inbreeding depression and offspring sex.     

Geographical variation in offspring size effects across generations

Offspring size is thought to strongly affect offspring fitness and many studies have shown strong offspring size/fitness relationships in marine and terrestrial organisms. This relationship is strongly mitigated by local environmental conditions and the optimal offspring size that mothers should produce will vary among different environments. It is assumed that offspring size will consistently affect the same traits among populations but this assumption has not been tested. Here I use a common garden experiment to examine the effects of offspring size on subsequent performance for the marine bryozoan Bugula neritina using larvae from two very different populations. The local conditions at one population (Williamstown) favour early reproduction whereas the other population (Pt. Wilson) favours early growth. Despite being placed in the same habitat, the effects of parental larval size were extremely variable and crossed generations. For larvae from Williamstown, parental larval size positively affected initial colony growth and larval size in the next generation. For larvae from the other population, parental larval size positively affected colony fecundity and negatively affected larval size in the next generation. Traditionally, exogenous factors have been viewed as the sole source of variation in offspring size/fitness relationship but these results show that endogenous factors (maternal source population) can also cause variation in this crucial relationship. It appears offspring size effects can be highly variable among populations and organisms can adapt to local conditions without changing the size of their offspring.     

The short-term and long-term effects of parental age in the bean weevil (Acanthoscelides obtectus)

We examined the influence of parental age on life history traits of their offspring in the lines of bean weevil that have evolved different rates of senescence. Measurements included preadult traits (egg size, embryonic developmental time, total preadult developmental time, preadult viability) and adult traits (body weight, total realized fecundity of females, first day of egg laying, early fecundity, late fecundity and longevity). The negative parental age effects were observed for all traits except for the early and total realized fecundity. We did not detect statistically significant line×parental age interactions for either preadult- or adult-survival, so offspring survival did not change with parental age after selection for early vs. late reproduction. It seems that selection acting on the quality of offspring produced by parents of different ages has not been responsible for the evolution of senescence in bean weevil. This revised version was published online in August 2006 with corrections to the Cover Date.     

Reproductive senescence and parental effects in an indeterminate grower

Reproductive senescence is the decrease of reproductive performance with increasing age and can potentially include trans‐generational effects as the offspring produced by old parents might have a lower fitness than those produced by young parents. This negative effect may be caused either by the age of the father, mother or the interaction between the ages of both parents. Using the common woodlouse Armadillidium vulgare, an indeterminate grower, as a biological model, we tested for the existence of a deleterious effect of parental age on fitness components. Contrary to previous findings reported from vertebrate studies, old parents produced both a higher number and larger offspring than young parents. However, their offspring had lower fitness components (by surviving less, producing a smaller number of clutches or not reproducing at all) than offspring born to young parents. Our findings strongly support the existence of trans‐generational senescence in woodlice and contradict the belief that old individuals in indeterminate growers contribute the most to recruitment and correspond thereby to the key life stage for population dynamics. Our work also provides rare evidence that the trans‐generational effect of senescence can be stronger than direct reproductive senescence in indeterminate growers.     

What happens to offspring when parents are inbred,old or had a poor start in life? Evidence for sex‐specific parental effects

Parental effects on offspring performance have been attributed to many factors such as parental age, size and condition. However, we know little about how these different parental characteristics interact to determine parental effects, or the extent to which their effect on offspring depends on either the sex of the parent or that of the offspring. Here we experimentally tested for effects of variation in parents’ early diet and inbreeding levels, as well as effects of parental age, and for potential interactive effects of these three factors on key aspects of offspring development in the mosquitofish (Gambusia holbrooki). Older mothers produced offspring that were significantly smaller at birth. This negative effect of maternal age on offspring size was still evident at maturation as older mothers had smaller daughters, but not smaller sons. The daughters of older mothers did, however, reach maturity sooner. Paternal age did not affect offspring body size, but it had a complex effect on their sons’ relative genital size. When initially raised on a food‐restricted diet, older fathers sired sons with relatively smaller genitalia, but when fathers were initially raised on a control diet their sons had relatively larger genitalia. The inbreeding status of mothers and fathers had no significant effects on any of the measured offspring traits. Our results indicate that the manifestation of parental effects can be complex. It can vary with both parent and offspring sex; can change over an offspring's life; and is sometimes evident as an interaction between different parental traits. Understanding this complexity will be important to predict the role of parental effects in adaptation.     

Heterozygosity at a single locus explains a large proportion of variation in two fitness‐related traits in great tits: a general or a local effect?

In natural populations, mating between relatives can have important fitness consequences due to the negative effects of reduced heterozygosity. Parental level of inbreeding or heterozygosity has been also found to influence the performance of offspring, via direct and indirect parental effects that are independent of the progeny own level of genetic diversity. In this study, we first analysed the effects of parental heterozygosity and relatedness (i.e. an estimate of offspring genetic diversity) on four traits related to offspring viability in great tits (Parus major) using 15 microsatellite markers. Second, we tested whether significant heterozygosity–fitness correlations (HFCs) were due to ‘local’ (i.e. linkage to genes influencing fitness) and/or ‘general’ (genome‐wide heterozygosity) effects. We found a significant negative relationship between parental genetic relatedness and hatching success, and maternal heterozygosity was positively associated with offspring body size. The characteristics of the studied populations (recent admixture, polygynous matings) together with the fact that we found evidence for identity disequilibrium across our set of neutral markers suggest that HFCs may have resulted from genome‐wide inbreeding depression. However, one locus (Ase18) had disproportionately large effects on the observed HFCs: heterozygosity at this locus had significant positive effects on hatching success and offspring size. It suggests that this marker may lie near to a functional locus under selection (i.e. a local effect) or, alternatively, heterozygosity at this locus might be correlated to heterozygosity across the genome due to the extensive ID found in our populations (i.e. a general effect). Collectively, our results lend support to both the general and local effect hypotheses and reinforce the view that HFCs lie on a continuum from inbreeding depression to those strictly due to linkage between marker loci and genes under selection.     

Fecundity selection theory: concepts and evidence

Fitness results from an optimal balance between survival, mating success and fecundity. The interactions between these three components of fitness vary depending on the selective context, from positive covariation between them, to antagonistic pleiotropic relationships when fitness increases in one reduce the fitness of others. Therefore, elucidating the routes through which selection shapes life history and phenotypic adaptations via these fitness components is of primary significance to understanding ecological and evolutionary dynamics. However, while the fitness components mediated by natural (survival) and sexual (mating success) selection have been debated extensively from most possible perspectives, fecundity selection remains considerably less studied. Here, we review the theoretical basis, evidence and implications of fecundity selection as a driver of sex‐specific adaptive evolution. Based on accumulating literature on the life‐history, phenotypic and ecological aspects of fecundity, we (i) suggest a re‐arrangement of the concepts of fecundity, whereby we coin the term ‘transient fecundity’ to refer to brood size per reproductive episode, while ‘annual’ and ‘lifetime fecundity’ should not be used interchangeably with ‘transient fecundity’ as they represent different life‐history parameters; (ii) provide a generalized re‐definition of the concept of fecundity selection as a mechanism that encompasses any traits that influence fecundity in any direction (from high to low) and in either sex; (iii) review the (macro)ecological basis of fecundity selection (e.g. ecological pressures that influence predictable spatial variation in fecundity); (iv) suggest that most ecological theories of fecundity selection should be tested in organisms other than birds; (v) argue that the longstanding fecundity selection hypothesis of female‐biased sexual size dimorphism (SSD) has gained inconsistent support, that strong fecundity selection does not necessarily drive female‐biased SSD, and that this form of SSD can be driven by other selective pressures; and (vi) discuss cases in which fecundity selection operates on males. This conceptual analysis of the theory of fecundity selection promises to help illuminate one of the central components of fitness and its contribution to adaptive evolution.     

Early reproductive investment,senescence and lifetime reproductive success in female Asian elephants

The evolutionary theory of senescence posits that as the probability of extrinsic mortality increases with age, selection should favour early‐life over late‐life reproduction. Studies on natural vertebrate populations show early reproduction may impair later‐life performance, but the consequences for lifetime fitness have rarely been determined, and little is known of whether similar patterns apply to mammals which typically live for several decades. We used a longitudinal dataset on Asian elephants (Elephas maximus) to investigate associations between early‐life reproduction and female age‐specific survival, fecundity and offspring survival to independence, as well as lifetime breeding success (lifetime number of calves produced). Females showed low fecundity following sexual maturity, followed by a rapid increase to a peak at age 19 and a subsequent decline. High early life reproductive output (before the peak of performance) was positively associated with subsequent age‐specific fecundity and offspring survival, but significantly impaired a female's own later‐life survival. Despite the negative effects of early reproduction on late‐life survival, early reproduction is under positive selection through a positive association with lifetime breeding success. Our results suggest a trade‐off between early reproduction and later survival which is maintained by strong selection for high early fecundity, and thus support the prediction from life history theory that high investment in reproductive success in early life is favoured by selection through lifetime fitness despite costs to later‐life survival. That maternal survival in elephants depends on previous reproductive investment also has implications for the success of (semi‐)captive breeding programmes of this endangered species.     

Maternal caloric restriction partially rescues the deleterious effects of advanced maternal age on offspring

While many studies have focused on the detrimental effects of advanced maternal age and harmful prenatal environments on progeny, little is known about the role of beneficial non‐Mendelian maternal inheritance on aging. Here, we report the effects of maternal age and maternal caloric restriction (CR) on the life span and health span of offspring for a clonal culture of the monogonont rotifer Brachionus manjavacas. Mothers on regimens of chronic CR (CCR) or intermittent fasting (IF) had increased life span compared with mothers fed ad libitum (AL). With increasing maternal age, life span and fecundity of female offspring of AL‐fed mothers decreased significantly and life span of male offspring was unchanged, whereas body size of both male and female offspring increased. Maternal CR partially rescued these effects, increasing the mean life span of AL‐fed female offspring but not male offspring and increasing the fecundity of AL‐fed female offspring compared with offspring of mothers of the same age. Both maternal CR regimens decreased male offspring body size, but only maternal IF decreased body size of female offspring, whereas maternal CCR caused a slight increase. Understanding the genetic and biochemical basis of these different maternal effects on aging may guide effective interventions to improve health span and life span.     

Rapid within‐ and transgenerational changes in thermal tolerance and fitness in variable thermal landscapes

Phenotypic plasticity may increase the performance and fitness and allow organisms to cope with variable environmental conditions. We studied within‐generation plasticity and transgenerational effects of thermal conditions on temperature tolerance and demographic parameters in Drosophila melanogaster. We employed a fully factorial design, in which both parental (P) and offspring generations (F1) were reared in a constant or a variable thermal environment. Thermal variability during ontogeny increased heat tolerance in P, but with demographic cost as this treatment resulted in substantially lower survival, fecundity, and net reproductive rate. The adverse effects of thermal variability (V) on demographic parameters were less drastic in flies from the F1, which exhibited higher net reproductive rates than their parents. These compensatory responses could not totally overcome the challenges of the thermally variable regime, contrasting with the offspring of flies raised in a constant temperature (C) that showed no reduction in fitness with thermal variation. Thus, the parental thermal environment had effects on thermal tolerance and demographic parameters in fruit fly. These results demonstrate how transgenerational effects of environmental conditions on heat tolerance, as well as their potential costs on other fitness components, can have a major impact on populations’ resilience to warming temperatures and more frequent thermal extremes.     

BIRTH‐ORDER DIFFERENCES CAN DRIVE NATURAL SELECTION ON AGING

Senescence—the deterioration of survival and reproductive capacity with increasing age—is generally held to be an evolutionary consequence of the declining strength of natural selection with increasing age. The diversity in rates of aging observed in nature suggests that the rate at which age‐specific selection weakens is determined by species‐specific ecological factors. We propose that, in iteroparous species, relationships between parental age, offspring birth order, and environment may affect selection on senescence. Later‐born siblings have, on average, older parents than do first borns. Offspring born to older parents may experience different environments in terms of family support or inherited resources, factors often mediated by competition from siblings. Thus, age‐specific selection on parents may change if the environment produces birth‐order related gradients in reproductive success. We use an age‐and‐stage structured population model to investigate the impact of sibling environmental inequality on the expected evolution of senescence. We show that accelerated senescence evolves when later‐born siblings are likely to experience an environment detrimental to lifetime reproduction. In general, sibling inequality is likely to be of particular importance for the evolution of senescence in species such as humans, where family interactions and resource inheritance have important roles in determining lifetime reproduction.     

Strain‐specific effects of parental age on offspring in Drosophila melanogaster

Maternal age is generally known to be negatively correlated with the lifespan of offspring in several animal models including yeast, rotifers, flies, and possibly in humans. However, several reports have shown positive effects of parental age on offspring lifespan. Thus, there was a need to investigate further the inconsistent results on the effect of parental age on lifespan. In this study, the effects of parental age on offspring fitness and lifespan were examined by using Drosophila melanogaster. The lifespan of offspring from old parents was significantly increased compared with that of the young counterparts in the Canton‐S (CS) strain but not in other D. melanogaster strains, such as Oregon‐R (OR) and w¹¹¹⁸. To find out why the lifespan is increased in the offspring from old parents in CS flies, fitness components that could modulate lifespan were examined in CS flies. Egg weight and body weight were reduced by parental aging and the offspring of old fathers or old mothers developed faster than that of the young. In addition, the offspring of old parents had increased resistance to oxidative and heat shock stresses. However, reproductive capacity, mating preference, and food intake were unaffected by parental aging. These results indicate that parental aging in CS strain D. melanogaster has beneficial effects on the lifespan and fitness of offspring. The presence of strain‐specific manner effects suggests that genetic background might be a significant factor in the parental age effect.     

Effects of parental light environment on growth and morphological responses of clonal offspring

• Environments experienced by parent ramets of clonal plants can potentially influence fitness of clonal offspring ramets. Such clonal parental effects may result from heritable epigenetic changes, such as DNA methylation, which can be removed by application of DNA de‐methylation agents such as 5‐azacytidine.
• To test whether parental shading effects occur via clonal generation and whether DNA methylation plays a role in such effects, parent plants of the clonal herb Alternanthera philoxeroides were first subjected to two levels of light intensity (high versus low) crossed with two levels of DNA de‐methylation (no or with de‐methylation by application of 5‐azacytidine), and then clonal offspring taken from each of these four types of parent plant were subjected to the same two light levels.
• Parental shading effects transmitted via clonal generation decreased growth and modified morphology of clonal offspring. Offspring responses were also influenced by DNA methylation level of parent plants. For clonal offspring growing under low light, parental shading effects on growth and morphology were always negative, irrespective of the parental de‐methylation treatment. For clonal offspring growing under high light, parental shading effects on offspring growth and morphology were negative when the parents were not treated with 5‐azacytidine, but neutral when they were treated with 5‐azacytidine.
• Overall, parental shading effects on clonal offspring performance of A. philoxeroides were found, and DNA methylation is likely to be involved in such effects. However, parental shading effects contributed little to the tolerance of clonal offspring to shading.

     

On the virtue of being the first born: the influence of date of birth on fitness in the mosquitofish, Gambusia affinis

We found in an earlier study that mosquitofish (Gambusia affinis and G. holbrooki) ceased reproduction in the late summer, long before the end of warm weather, stored fat, then utilized reserves to survive the winter and initiate reproduction the following spring. We hypothesized that this pattern of fat utilization was a life history adaptation that enabled the fish to acquire food resources in the autumn then allocate them to reproduction the following spring when the fitness of the young would be greater. Here we evaluate one aspect of this hypothesis by evaluating the probability of survival to maturity and fecundity of young as a function of date of birth. We placed cohorts comprising eight to ten litters of young born early‐, mid‐ or late in the reproductive season in replicate field enclosures. The entire experiment was repeated in two different years. Early‐born young had a significantly higher probability of survival to maturity but did not differ in fecundity relative to the last cohort of the season. Early‐born young also attained maturity early enough to reproduce in their year of birth while late‐born young had to overwinter before reproduction. The fitness consequences to the mother of either producing one more litter of young at the end of the season, versus instead storing fat and reproducing the following spring are not as determinate as are the effects of date of birth on offspring fitness. Females most often gain fitness by not producing one last litter and instead over‐wintering. If, however, the overwinter survival of offspring is not influenced by their size at the end of the season, then a female's fitness could be enhanced by producing one more litter late in the season. If instead the probability of overwinter survival is strongly influenced by the size of offspring at the end of the season, then our results suggest that a female gains more by deferring reproduction and storing for overwinter survival and reproduction the following spring.     

Parent–offspring resemblance in colony-specific adult survival of cliff swallows

Survival is a key component of fitness. Species that occupy discrete breeding colonies with different characteristics are often exposed to varying costs and benefits associated with group size or environmental conditions, and survival is an integrative net measure of these effects. We investigated the extent to which survival probability of adult (≥1-year old) cliff swallows (Petrochelidon pyrrhonota) occupying different colonies resembled that of their parental cohort and thus whether the natal colony had long-term effects on individuals. Individuals were cross-fostered between colonies soon after hatching and their presence as breeders monitored at colonies in the western Nebraska study area for the subsequent decade. Colony-specific adult survival probabilities of offspring born and reared in the same colony, and those cross-fostered away from their natal colony soon after birth, were positively and significantly related to subsequent adult survival of the parental cohort from the natal colony. This result held when controlling for the effect of natal colony size and the age composition of the parental cohort. In contrast, colony-specific adult survival of offspring cross-fostered to a site was unrelated to that of their foster parent cohort or to the cohort of non-fostered offspring with whom they were reared. Adult survival at a colony varied inversely with fecundity, as measured by mean brood size, providing evidence for a survival–fecundity trade-off in this species. The results suggest some heritable variation in adult survival, likely maintained by negative correlations between fitness components. The study provides additional evidence that colonies represent non-random collections of individuals.     

Larval population density affects female weight and fecundity in the dung beetle Aphodius ater

1. Competition for food at high densities during larval development leads to reduced adult weight in the northern temperate dung beetle Aphodius ater. 2. Analysis of female beetles caught in the field showed that numbers of eggs and total egg load per female were correlated positively with beetle size. 3. Female beetles reared at different population densities during larval development in the laboratory were analysed with regard to their lifetime fecundity and reproductive lifespan. 4. High population densities during development had a negative influence on the number of eggs per female and on reproductive lifespan. Lifetime fecundity was correlated positively with female weight. 5. It was concluded that competition during larval development in the first generation of offspring will result in a lower number of offspring in the second generation in Aphodius ater, and thereby reduce parental fitness.     

The pay‐offs of maternal care increase as offspring develop,favouring extended provisioning in an egg‐feeding frog

Offspring quantity and quality are components of parental fitness that cannot be maximized simultaneously. When the benefits of investing in offspring quality decline, parents are expected to shift investment towards offspring quantity (other reproductive opportunities). Even when mothers retain complete control of resource allocation, offspring control whether to allocate investment to growth or development towards independence, and this shared control may generate parent–offspring conflict over the duration of care. We examined these predictions by, in a captive colony, experimentally removing tadpoles of the strawberry poison frog (Oophaga pumilio) from the mothers that provision them with trophic eggs throughout development. Tadpoles removed from their mothers were no less likely to survive to nutritional independence (i.e. through metamorphosis) than were those that remained with their mothers, but these offspring were smaller at metamorphosis and were less likely to survive to reach adult size, even though they were fed ad libitum. Tadpoles that remained with their mothers developed more slowly than those not receiving care, a pattern that might suggest that offspring extracted more care than was in mothers’ best interests. However, the fitness returns of providing care increased with offspring development, suggesting that mothers would be best off continuing care until tadpoles initiated metamorphosis. Although the benefits of parental investment in offspring quality are often thought to asymptote at high levels, driving parent–offspring conflict over weaning, this assumption may not hold over natural ranges of investment, with selection on both parents and offspring favouring extended durations of parental care.