Mixed sex allocation strategies in a polytocous mammal,the house mouse (<Emphasis Type="Italic">Mus musculus</Emphasis>)

The issue of adaptive adjustment of offspring sex ratio (proportion of male births) in polytocous mammals, producing several offspring per litter, is controversial because females of these species can maximize their fitness mainly by adjusting offspring number. To address this issue, we examined the effect of maternal condition at mating, experimentally decreased by pre-mating food restriction, on the sex ratio variation in 137 female mice. We tested two basic sex allocation hypotheses plausible for polytocous mammals: (1) the Myers hypothesis, predicting that cheaper sex should be favored in poor environmental conditions to maximize offspring number; and (2) the Williams hypothesis, predicting maximum fitness returns by adjusting size- and sex-specific composition of the litter according to the maternal condition. The food-restricted mothers produced larger litters with a higher proportion of cheaper daughters than the control mothers. By contrast, the control mothers optimized size and sex composition of the litter according to their weight at mating. In addition, the offspring of the food-restricted mothers suffered less from pre-weaning mortality than those of the control mothers. Therefore, when comparing the groups, the Myers hypothesis had a general significance while the Williams hypothesis was plausible only for the control mothers. Furthermore, some of the food-restricted mothers partly coped with the pre-mating food restriction and increased the proportion of sons in the litter with the increasing maternal weight loss (during the period of food restriction). The sex ratio variation was thus a result of three sex allocation strategies depending on the maternal condition at mating.     

Factors influencing length of maternal care in brown bears (<Emphasis Type="Italic">Ursus arctos</Emphasis>) and its effect on offspring

Length of maternal care, i.e. the interval between successfully raised litters, is the most important factor explaining the variation in reproductive rate among brown-bear (Ursus arctos) populations. In this paper, we examine the variation in length of maternal care in radio-marked brown bears and its effect on their offspring in northern Sweden. Young stayed with their mothers for 1.4–1.5 or 2.4–2.5 (in one case 3.5) years and were weaned with body masses varying from 17 to 69 kg. The probability of yearling litters staying with their mother for a 2nd year increased with decreasing yearling body mass, and was higher for litters with two offspring than for litters with one or three to four offspring. Staying with their mothers for a 2nd year had a positive effect on mass gain in yearlings and this effect was more pronounced in litters with two than three to four offspring. Body mass of 2-year-olds was not related to age of weaning, suggesting that keeping offspring for an additional year mainly compensated for low yearling body mass. If large offspring body mass positively affects later offspring survival and reproduction, mothers may be able to optimize the length of maternal care according to the litter size and the size of their yearlings.Communicated by F. Trillmich     

Effects of maternal nutrition on post-weaning growth in two North American rodents

The Trivers-Willard hypothesis of sex-biased maternal investment in response to fluctuations in resource availability has provided a theoretical foundation for research on maternal investment for more than two decades. Their hypothesis holds that mothers in poor condition as a result of poor resource availability should bias parental investment towards offspring with the highest probability of reproducing. In the polygynous mating system of mammals where males compete for breeding access to females, the hypothesis predicts investment favoring females. Although data from many systems have supported this hypothesis, other systems do not follow the predicted patterns and have resulted in various alternative hypotheses. The present research was designed to test whether differences in body condition of young reared by nutritionally stressed dams relative to young reared by unstressed dams were maintained into adulthood, one of the fundamental assumptions underlying the Trivers-Willard hypothesis. Post-weaning growth in eastern woodrats (Neotoma floridana) and northern grasshopper mice (Onychomys leucogaster) was examined relative to maternal nutritional plane. Individuals from undernourished dams were lighter than their unrestricted counterparts at weaning but no difference was evident by the time they had reached adult size. Failure to maintain body condition differences into adulthood violates one of the assumptions essential for application of the Trivers-Willard hypothesis of maternal investment patterns. Although the Trivers-Willard model proposed that natural selection favors differential investment in the sexes over the entire course of parental investment, evidence from this and other studies suggests that the Trivers-Willard hypothesis might not be appropriate to address maternal investment questions in postnatally malnourished dams, but instead should be restricted to systems concerned with prenatal maternal condition or resource availability. Received: 22 February 1995/Accepted after revision: 30 December 1995     

Climate and maternal effects modify sex ratios in a weakly dimorphic marsupial

There is growing evidence that the sex ratios of wild vertebrate populations are determined by mechanisms that are directly influenced by environmental characteristics. The Trivers–Willard (TWH) and extrinsic modification (EMH) hypotheses postulate differing determinants of mammalian offspring sex ratios. TWH states that mothers allocate resources according to their current condition and sex-specific offspring costs. EMH states that environmental forces that affect maternal condition determine offspring sex ratios, independently of maternal tactics of sex-biased allocation. We statistically assessed support for each of these hypotheses using long-term life histories of the allied rock-wallaby, Petrogale assimilis; a continuously breeding, polygynous, weakly dimorphic marsupial. We showed that birth sex ratios were equal and independent of maternal and environmental conditions. However, secondary sex ratios were male-biased under good environmental conditions and for high quality mothers or mothers in good condition. Sex differences in offspring survival contributed to these biases: (1) environmental conditions strongly influenced survival to pouch emergence (in support of EMH) and (2) maternal quality affected survival to the end of maternal care (in support of TWH). Environmental effects on survival were more important than maternal factors over the entire period of maternal care and contributed most to male-biased sex ratios at pouch emergence. In contrast, maternal mass was the best predictor of sex ratios at the end of maternal care—the life history stage where offspring body mass differed between the sexes.     

Aging and improving reproductive success in horses: declining residual reproductive value or just older and wiser?

In many mammalian species, female success in raising offspring improves as they age. The residual reproductive value hypothesis predicts that each individual offspring will be more valuable to the mother as she ages because there is less conflict between the current and potential future offspring. Therefore, as mothers age, their investment into individual offspring should increase. Empirical evidence for an influence of declining residual reproductive value on maternal investment is unconvincing. Older mothers may not invest more, but may be more successful due to greater experience, allowing them to target their investment more appropriately (targeted reproductive effort hypothesis). Most studies do not preclude either hypothesis. Mare age significantly influenced maternal investment in feral horses living on the North Island of New Zealand. Older mares, that were more successful at raising foals, were more protective for the first 20 days of life, but less diligent thereafter. Total maternal input by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival and a similar pattern was observed in mares that had lost a foal in the previous year. In addition, older mothers were more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring. Therefore, older mothers seem to become more successful by targeting their investment better due to experience, not by investing more in their offspring. Received: 22 September 1999 / Received in revised form: 30 October 1999 / Accepted: 6 November 1999     

Litter reductions reveal a trade-off between offspring size and number in brown bears

Experimental manipulations have revealed positive effects of litter reduction on offspring mass in small mammals, but little is known about this trade-off in large mammals. We examined the determinants of natural litter size variation and quantified the effects of litter size, maternal characteristics, and litter composition on yearling mass using 24?years of data on marked brown bears (Ursus arctos) in Sweden. Infanticide by adult males is a major cause of cub-of-the-year mortality, leading to litter size reductions. Litter size (n?=?265) at den emergence ranged from one to four cubs (average, 2.7) and increased with maternal age. Litter size, however, appeared independent of maternal size, population density, interlitter interval, study area, or previous litter sex ratio. Yearling body mass increased with maternal body size but was independent of litter sex ratio. Litter size and yearling mass were negatively correlated, mostly because singletons were about 30?% heavier than yearlings from litters of two to four cubs. In reduced litters, survivors were on average 8?% heavier as yearlings than individuals from intact litters, suggesting that sibling competition reduces growth. Trade-offs between litter size and yearling mass in bears appear similar in magnitude to those found in small mammals.     

Contradictory findings in studies of sex ratio variation in roe deer (Capreolus capreolus)

Patterns of sex ratio variation and maternal investment reported in the literature are often inconsistent. This could be due to intra- and inter-specific variation in social systems, but may also be a result of the a posteriori nature of much of this type of analysis or the testing of models which are inappropriate. Two recent papers reported directly opposed results concerning variation in offspring sex ratios in relation to maternal condition in roe deer, interpreting the results as support for the Trivers and Willard model and for the local resource competition hypothesis, respectively. In this paper, we present data on offspring sex ratios and early juvenile body weight from two long-term studies of this species to test predictions arising from these two models concerning sex biases in litter composition and maternal care. First, we observed no consistent pattern of sex differences in an index of weaning weight or body weight at 1 month old in either population, indicating a lack of sex bias in maternal care. However, in one population, higher maternal body weight was associated with higher juvenile body weight of daughters, but not of sons. Secondly, we found a negative, but not statistically significant, relationship between maternal body weight and litter sex ratio such that heavier females tended to produce more daughters and lighter females to produce more sons. These results indicate that roe females which have additional investment potential available do not invest it in sons, as predicted by the Trivers and Willard model. Our results may provide some support that roe deer are subject to local resource competition acting at the level of the individual mother; however, the fact that particular trends in sex ratio data can be explained in functional terms provides no indication that they are actually adaptive. Received: 9 December 1997 / Accepted after revision: 11 November 1998     

Maternal investment, sex-differential prospects, and the sex ratio in wild house mice

In a population of first-generation offspring from wild-caught house mice (Mus musculus domesticus), previous evidence suggested that male fitness is more strongly affected by an increase in body weight than female fitness. This paper shows that in these mice the young are weaned at heavier weights the smaller the litter and the better the maternal body condition. These effects persisted into adulthood and were less pronounced in female young. However, contrary to expectation from conventional sex ratio theory, maternal condition and litter size had no detectable effect on sex ratios. Also, litter size did not affect sex ratios in two populations of laboratory-kept, wild-caught western (M. m. domesticus) and eastern house mice (M. m. musculus). Wild house mice, therefore, appear not to adaptively manipulate the sex ratio of offspring. It is argued that this absence of sex ratio trends might not be maladaptive, but rather that models currently used to predict sex ratio trends in rodents may not be valid. Received: 13 March 1997 / Accepted after revision: 9 August 1997     

Female-biased sex allocation of offspring by an <Emphasis Type="Italic">Apodemus</Emphasis> mouse in an unstable environment

We investigated the effects of population fluctuation on the offspring’s sex allocation by a weakly polygynous mouse, Apodemus argenteus, for 3 years. In acorn-poor seasons, heavier mothers invested more in sons, and lighter mothers invested more in daughters. In acorn-rich seasons, heavier mothers invested more in daughters, and lighter mothers invested more in sons. Maternal body condition and litter size affected the sex allocation. Furthermore, there was a maternal investment trade-off between a son’s birth mass and the number of daughters. Based upon the effect of population fluctuation on the lifetime reproductive success of each sex, we proposed the new “safe bet hypothesis”. This hypothesis predicts that frequent and unpredictable change in female distribution, which is often caused by abrupt fall in food condition, favors female-biased maternal investment to offspring by polygynous mammals and is applicable to many small mammals inhabiting in unstable environments.     

Lactation performance of guinea pigs (Cavia porcellus) does not respond to experimental manipulation of pup demands

To ensure proper development of young, mothers should react to offspring signals of need. Studies of such parent-offspring interaction often manipulated litter size to measure effects of changed offspring food demand. We used the extreme precociality of guinea pig (Cavia porcellus) pups to increase offspring demand without changing any other litter characteristic. As pups contribute to their own energy demands from birth by independent feeding, their food demand can be increased by withholding solid food early in lactation. We studied whether mothers reacted to increased food demand of offspring by enhanced parental care, especially by changes in the pattern of milk production and nursing. Pups deprived of solid food early in lactation grew more slowly and were in poorer body condition than pups in control litters, even after the former had access to solid food in late lactation. Mothers of deprived young reacted to offspring long-term need by maintaining nursing behaviour for longer than control mothers. However, this change in behaviour did not occur early in lactation when pup short-term need was greatest nor did it result in increased milk transfer at any time. Energy allocation of mothers measured by changes in their food intake, maternal body reserves, and milk production stayed the same whether offspring had early access to solid food or not. Thus mothers did not increase energy allocation to pups even though they apparently had information about the pups' poor state.     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Intergenerational effects of climate generate cohort variation in lizard reproductive performance

An evaluation of the link between climate and population dynamics requires understanding of climate effects both within and across generations. In ectothermic vertebrates, demographic responses to climate changes should crucially depend on balancing needs for heat and water. Here, we studied how temperature and rainfall regimes experienced before and during adulthood influenced reproductive performances (litter size, offspring size, and survival) in a natural population of the live-bearing common lizard, Lacerta vivipara, monitored continuously from 1989 to 2004. Rainfall regime, but not temperature, had both immediate and delayed effects on these reproductive performances. Rainfall during the first month of life was positively correlated with juvenile survival. Females experiencing more rainfall during gestation produced smaller neonates that showed greater survival when controlling for the positive effect of body size on survival. Furthermore, females that experienced heavier rainfall when in utero produced fewer but longer neonates during adulthood. These demographic effects of rainfall on adult reproductive traits may come from maternal effects of climate conditions and/or from delayed effects of rainfall on the environment experienced early in life. Irrespective of the precise mechanism, however, this study provides evidence of intergenerational climate effects in natural populations of an ectothermic vertebrate.     

Sex-specific preweaning maternal care in reindeer<Emphasis Type="Italic"> (Rangifer tarandus t.)</Emphasis>

Caribou (Rangifer tarandus) have been used as an example of a polygynous species in which one could expect females to manipulate their investments in sons and daughters based on their physical condition. We investigated whether the effect of maternal condition, measured as female body mass, on preweaning maternal care measured as calf mass in mid-July, calf mass in mid-November and calf mass gain (growth between mid-July and mid-November), was sex-biased in reindeer. We analysed separately young (h 4 years), prime age (5-10 years) and old (>10 years) mothers. Consistently, preweaning maternal care increased with increasing mother quality, but maternal condition did not influence whether a female produced a son or a daughter. Moreover, regardless of whether a female was young, prime age or old, there was no conclusive evidence of female quality-related, sex-biased preweaning maternal care in the study population. Our results indicate that maternal expenditure may be of equal magnitude in male and female offspring, regardless of the mothers' physical condition, and may support the hypothesis that a high level of maternal expenditure limits sex-biased maternal care.     

Costs and benefits of communal rearing to female prairie voles (<Emphasis Type="Italic">Microtus ochrogaster</Emphasis>)

Communal rearing of offspring may help mothers maximize their investment in offspring at a reduced cost to their own bodily condition, thus maximizing their potential for reproductive success. The objective of this study was to quantify the costs and benefits of communal rearing to prairie vole (Microtus ochrogaster) pups and mothers. Mothers were assigned to one of three social units: solitary mothers, singularly breeding groups (i.e. one mother and one non-reproductive sister) and plurally breeding groups (i.e. two lactating sisters). For each type of social unit, some replicates were provided with food ad libitum, while others were provided with limited food. The body mass of focal mothers (i.e. the first mother to produce a litter) was a significant predictor of pup growth. Regardless of food availability, litters of focal mothers in plurally breeding groups gained more weight than litters reared by solitary mothers. Pups reared in singularly breeding groups were intermediate in weight gain, but did not gain significantly more weight than solitary offspring. There was no difference in the body mass of focal mothers from each type of social unit, regardless of food availability. Within plurally breeding groups, the weight gain of the two litters and body mass of focal and second mothers did not differ. However, focal mothers from plurally breeding groups nursed fewer pups than solitary mothers and also fewer pups than their nestmates when food was limited. Our results suggest that plural breeding results in greater fitness to mothers than solitary and singular breeding.Communicated by E. Korpimäki     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Maternal care in house mice

Summary This paper analyzes the flexibility of maternal care in wild house mice (Mus domesticus) under different reproductive conditions in the laboratory. All maternal activities were both qualitatively and quantitatively analyzed over a period of 28 days after birth of a litter. The standard behavior of a lactating house mouse with 7–8 young can be described as follows: During days 1–16 the offspring fully depend on the mother for nutrition. Due to rapid growth of the litter, the energetic demands of lactation reach a peak for the female during days 13–16. During days 17–22, the weaning period, the young begin to eat solid food. This period is characterized by behaviors that indicate different interests of the mother and offspring, and thus the existence of a parent-offspring conflict sensu Trivers (1974). Resting alone and remaining far from the litter indicate the female's interest in avoiding the offspring's demands, which are expressed in frequent attempts to initiate sucking. There is no aggression towards the young during weaning. House mice are weaned at 23 days. The relationship between mother and young appears free of conflict after weaning. Nursing is replaced by resting with body contact, but the offspring do not try to suck. The following results suggest that during the weaning period the offspring do not get more milk than corresponds to the maternal optimum—despite their frequent sucking attempts:(a) When the mother is simultaneously lactating and pregnant, offspring are smaller at weaning than under standard conditions. (b) Small litters are weaned earlier than large ones. Despite a longer nursing period, offspring from large litters are lighter at weaning than those from small ones. (c) Under high energy demand, as after postpartum mating and with large litters, females wean their young at a body weight which corresponds to the earliest physiologically possible state of independence.Parity of the female has no effect on maternal activities, nor has the presence of the father. In the latter case, however, offspring are less often left alone and unprotected.Females seem to adjust their investment according to the body weight of the progeny by delaying or advancing the date of weaning (Table 2). This behavior allows the production of the largest possible number of offspring that can be raised to a minimal physiological threshold corresponding to a body weight of approximately 9 g. Such flexibility in parental care may enhance maternal fitness under different and unpredictable environmental conditions.     

Maternal response to neonatal sibling conflict in the spotted hyena, <Emphasis Type="Italic">Crocuta crocuta</Emphasis>

Among spotted hyenas, Crocuta crocuta, neonatal aggression in twins is a well-known phenomenon and serves to establish intra-litter dominance soon after birth. As the stronger more aggressive cub presumably attains dominance over its twin, intra-litter dominance presents mothers with an ideal opportunity to assess individual cub fitness and, thereafter, to selectively favor one cub over the other. This study quantified maternal response to sibling conflict in 26 sets of twins born to 16 wild-living females to determine whether mothers of different social ranks exhibited favoritism towards sons or daughters, or in the case of same-sex twins, the dominant or subordinate cub. Maternal response to sibling conflict did not vary with litter sex composition, suggesting that mothers do not favor offspring of one sex over the other. All mothers intervened when their cubs fought and sometimes punished their cubs. Higher-ranking mothers more often punished both cubs, while lower-ranking mothers were more selective and punished the dominant cub. Where sibling aggression was most extreme, rather than favor the dominant sibling, mothers of all ranks made concessions to the subordinate cub that included procuring private nursing bouts for the subordinate and temporarily housing twins in separate dens, presumably to decrease sibling conflict. These findings represent a complex example of parent–offspring conflict and support both the insurance cub hypothesis and resource tracking hypothesis that mothers endeavor to keep all offspring alive for as long as possible in the event that the dominant sibling dies or that resources provide for the rearing of twins.     

Do female roe deer in good condition produce more sons than daughters

In polygynous roe deer Capreolus capreolus, males are only slightly heavier than females and the overall sex ratio at birth is close to unity. We studied offspring sex ratio and litter size (range 1–4, n = 74) of culled females, in utero, which provided an opportunity to examine responses of sex ratio to maternal condition. Male embryos were heavier than their sisters, and male fawns (9 months old) heavier than female fawns, suggesting a higher growth rate in males. There was no evidence for differential mortality between the sexes from birth to 9 months old. Heavier adult females produced larger embryos than lighter, or primiparous females. The overall sex ratio of embryos did not differ from unity, but adult does had more male embryos (55%) than primiparous does (32%), and the proportion of male embryos in a litter increased with the mother's body mass. Litter size also tended to increase with maternal age and body mass. We argue that this pattern reflects adaptive variation in offspring sex ratio.     

Benefits and costs of earwig (Forficula auricularia) family life

The evolution of parental care and family group formation critically depends on offspring survival benefits and parental fecundity costs of care under given ecological conditions. Investigations of the functional significance of care in insect species that exhibit facultative parental care have been relatively rare but may be of particular interest for better understanding of benefit and cost schedules at an early evolutionary stage. In this study, aspects of benefits and costs of care were addressed in the sub-social European earwig (Forficula auricularia; Dermaptera: Forficulidae) by manipulating the presence of tending mothers and brood size in a fully crossed experimental design. Larvae growing in broods tended by their mother or of reduced size showed a higher survival probability than larvae growing in untended or large broods, as predicted if maternal care is beneficial and shaped by a trade-off between number and quality of offspring. Analysis of patterns of food consumption and developmental time further suggested that the benefit of maternal attendance is mediated by the maternal provisioning of food, while the quality–quantity trade-off seemed to be driven by sibling rivalry. Further, tending mothers delayed the production of a second clutch, indicating a potential cost of care in terms of lifetime fecundity. This study experimentally shows benefits and potential costs of maternal care and family group formation in the European earwig. More detailed behavioural experiments will be required to fully understand how behavioural interactions among family members mediate these reproductive outcomes.     

Sex differences in the consequences of maternal loss in a long-lived mammal, the red deer (Cervus elaphus)

In several primates, the presence of mothers affects the growth, survival and reproduction of their offspring, but similar effects have not yet been demonstrated in ungulates. Here, we investigate the effects of the mother’s presence in a population of red deer (Cervus elaphus) on the Isle of Rum, Scotland, which is the subject of a long-term, individual-based study. We compared measures of performance including antler growth in young males and age at first reproduction in females and survival of deer with mothers still alive against those that have lost their mothers (orphans). We show that orphaning both before and after weaning increases the risk of a natural death for both sexes. For males, no maternal benefit was detectable past 24 months of age while, for females, post-weaning benefits continued throughout life. Orphaning resulted in compromised male physical condition as measured by a reduced probability of growing antlers by 16 months of age while no evidence for compromised reproduction was found in females. These results support assertions that post-weaning maternal associations affect the development and survival of offspring.