Genetic conflicts, intrinsic male fertility, and ejaculate investment

Few aspects of biology are linked to so many evolutionary conflicts as sperm production and fertilization. Segregation distortion and maternal inheritance of cytoplasmic genes, causing maladapted males, are common sources of variation in the competitive ability of sperm, leading males to vary in their intrinsic fertility. Here, I theoretically analyze the effect of such variation in male intrinsic fertility on ejaculate investment. The model reveals that with increasing variation in male fertility, males should overall spend less resources on their ejaculates. Furthermore, if males differing in intrinsic fertility are able to invest differently in sperm production, there are two contrasting outcomes. Typically, less fertile males should invest more. However, if female mating frequency is relatively low and differences between males relatively large, the most common male genotype should invest more. These results have important consequences both for the understanding of sperm competition strategies as well as for the evolution of female polyandry and female mating preferences.     

Placentation and maternal investment in mammals

The mammalian placenta exhibits striking interspecific morphological variation, yet the implications of such diversity for reproductive strategies and fetal development remain obscure. More invasive hemochorial placentas, in which fetal tissues directly contact the maternal blood supply, are believed to facilitate nutrient transfer, resulting in higher fetal growth rates, and to be a state of relative fetal advantage in the evolution of maternal-offspring conflict. The extent of interdigitation between maternal and fetal tissues has received less attention than invasiveness but is also potentially important because it influences the surface area for exchange. We show that although increased placental invasiveness and interdigitation are both associated with shorter gestations, interdigitation is the key variable. Gestation times associated with highly interdigitated labyrinthine placentas are 44% of those associated with less interdigitated villous and trabecular placentas. There is, however, no relationship between placental traits and neonatal body and brain size. Hence, species with more interdigitated placentas produce neonates of similar body and brain size but in less than half the time. We suggest that the effects of placental interdigitation on growth rates and the way that these are traded off against gestation length may be promising avenues for understanding the evolutionary dynamics of parent-offspring conflict.     

Growth, weaning and maternal investment from a comparative perspective

The process of weaning is related to a critical or threshold body weight attained by offspring among large-bodied mammals; the anthropoid primates, ungulates and pinnipeds. While weaning weight was allometrically related to maternal weight in interspecific comparisons, it was isometrically related to neonatal weight. When a neonate had grown to four times its birth weight, it was weaned. Differences between taxonomic groups were found only among the fasting phocids, where weanlings attained a lower, but proportional, weight. The duration of lactation was only weakly allometrically related to maternal or neonatal weight, and varied between individuals intraspecifically as a function of maternal condition. The time to weaning appears to be ecologically sensitive rather than to reflect interspecific life-history variation, in that, irrespective of the time to weaning, similar proportional weights appear to be attained. Interspecific similarities in threshold weaning weights are suggested to result from constraints on maternal abilities to meet energetic requirements of offspring through lactation after infants attain a threshold weight.     

Nutrition, fertility and steady-state population dynamics in a pre-industrial community in Penrith, northern England

The effect of nutrition on fertility and its contribution thereby to population dynamics are assessed in three social groups (elite, tradesmen and subsistence) in a marginal, pre-industrial population in northern England. This community was particularly susceptible to fluctuations in the price of grains, which formed their basic foodstuff. The subsistence class, who formed the largest part of the population, had low levels of fertility and small family sizes, but women from all social groups had a characteristic and marked subfecundity in the early part of their reproductive lives. The health and nutrition of the mother during pregnancy was the most important factor in determining fertility and neonatal mortality. Inadequate nutrition had many subtle effects on reproduction which interacted to produce a complex web of events. A population boom occurred during the second half of the 18th century; fertility did not change but there was a marked improvement in infant mortality and it is suggested that the steadily improving nutritional standards of the population, particularly during crucial periods in pregnancy (i.e. the last trimester), probably made the biggest contribution to the improvement in infant mortality and so was probably the major factor in triggering the boom.     

Changes in maternal investment in eggs can affect population dynamics

The way that mothers provision their offspring can have important consequences for their offspring's performance throughout life. Models suggest that maternally induced variation in life histories may have large population dynamical effects, even perhaps driving cycles such as those seen in forest Lepidoptera. The evidence for large maternal influences on population dynamics is unconvincing, principally because of the difficulty of conducting experiments at both the individual and population level. In the soil mite, Sancassania berlesei, we show that there is a trade-off between a female's fecundity and the per-egg provisioning of protein. The mother's position on this trade-off depends on her current food availability and her age. Populations initiated with 250 eggs of different mean sizes showed significant differences in the population dynamics, converging only after three generations. Differences in the growth, maturation and fecundity of the initial cohort caused differences in the competitive environment for the next generation, which, in turn, created differences in their growth and reproduction. Maternal effects in one generation can therefore lead to population dynamical consequences over many generations. Where animals live in environments that are temporally variable, we conjecture that maternal effects could result in long-term dynamical effects.     

Misregulation of phosphoinositides in Arabidopsis thaliana decreases pollen hydration and maternal fertility

Phosphoinositides are important lipids involved in membrane identity, vesicle trafficking, and intracellular signaling. In recent years, phosphoinositides have been shown to play a critical role in polarized secretion in plants, as perturbations of phosphoinositide metabolism, through loss of function mutants, result in defects in root hair elongation and pollen tube growth, where polarized secretion occurs rapidly. In the Brassicaceae, responses of stigmatic papillae to compatible pollen are also thought to involve highly regulated secretory events to facilitate pollen hydration and penetration of the pollen tube through the stigmatic surface. We therefore sought to analyze the female sporophyte fertility of the root hair defective4-1 mutant and the PI 4-kinase β1/β2 double mutant, which differentially affect phosphatidylinositol-4-phosphate (PI4P) levels. Stigmas from both mutants supported slower rates of pollen grain hydration, and the fecundity of these mutants was also diminished as a result of failed pollination events. This study therefore concludes that PI4P is integral to appropriate pistil responses to compatible pollen.     

Survival costs of reproduction predict age-dependent variation in maternal investment

Life-history theory predicts that older females will increase reproductive effort through increased fecundity. Unless offspring survival is density dependent or female size constrains offspring size, theory does not predict variation in offspring size. However, empirical data suggest that females of differing age or condition produce offspring of different sizes. We used a dynamic state-variable model to determine when variable offspring sizes can be explained by an interaction between female age, female state and survival costs of reproduction. We found that when costs depend on fecundity, young females with surplus state increase offspring size and reduce number to minimize fitness penalties. When costs depend on total reproductive effort, only older females increase offspring size. Young females produce small offspring, because decreasing offspring size is less expensive than number, as fitness from offspring investment is nonlinear. Finally, allocation patterns are relatively stable when older females are better at acquiring food and are therefore in better condition. Our approach revealed an interaction between female state, age and survival costs, providing a novel explanation for observed variation in reproductive traits.     

Sex-biased maternal investment in voles: importance of environmental conditions

Adaptive bias in sex allocation is traditionally proposed to be related to the condition of mothers as well as to the unequal fitness values of produced sexes. A positive relationship between mother condition and investment into male offspring is often predicted. This relationship was also recently found to depend on environmental conditions. We studied these causalities experimentally using a design where winter food supply was manipulated in eight outdoor-enclosed populations of field voles Microtus agrestis. At the beginning of the breeding season in spring, food-supplemented mothers seemed to be in a similar condition, measured as body mass, head width, body condition index and parasite load (blood parasite Trypanosoma), to non-supplemented mothers. Food supplements affected neither the litter size, the reproductive effort of mothers, nor the litter sex ratios at birth. However, food supplementation significantly increased the birth size of male offspring and improved their condition, as indicated by reduced parasite loads (intestinal Eimeria). Interestingly, mothers in good body condition produced larger male offspring only when environmental conditions were improved by food supplements. Although the adaptiveness of variation in mammalian sex ratios is still questionable, our study indicates that mothers in good condition bias their investment towards male offspring, but only when environmental conditions are favourable.     

Unequal maternal investment in offspring quality in relation to predation risk

Equal investment within broods does not always maximize parental reproductive value if the reproductive value of some of the young is low. We examined maternal investment in terms of offspring size in relation to the prospects of survival from predation within broods of the shield bug Elasmucha ferrugata Fabr. (Heteroptera; Acanthosomatidae). Shield bug females guard eggs and first instar nymphs against invertebrate predators by covering the clutch with their body and by behaving aggressively towards their enemies. Survival of eggs was not possible without maternal care. When females were allowed to guard their brood, eggs at the periphery were more vulnerable to predators than eggs at the centre. We found that females laid significantly larger eggs in the safest, central part of the clutch. There seems to be an advantage of large nymph size, since when nymphs were reared separately with low food resources, the larger ones were more likely to survive. Larger nymphs were also more likely to push themselves to the safest, central part of the clutch. Females seem to allocate their resources more to the offspring with the highest probability of avoiding predation. Thus our study supports unequal maternal investment within broods of E.ferrugata.     

The effect of paternal investment on female fertility intention in South Korea

Life history theory views reproduction as an outcome of resource allocation. The allocation of resources such as parental investments of time, energy and material resources involves trade-offs between number of offspring and timing of reproduction. Within the framework of mammalian parental investment, the outstanding feature of human reproduction is the high level of paternal care. Although empirical evidence suggests that human paternal investment may have evolved as a reproductive strategy to reduce infant and child mortality rates, the effects of actual paternal investment, including allocating time to child care, on female reproductive decisions have received relatively little attention. We examined the trade-off from two perspectives using a representative sample of married South Korean women aged 20–44 in 2005 (n=977). First, paternal investment in domestic labor, including child care and housework, was expected to be associated with women's preference regarding future reproduction. Second, relative paternal investment was expected to increase women's preference for future reproduction, especially among employed women. We found that increased paternal investment in child care and housework remarkably enhanced women's intention to have a second child, especially among employed women. In addition, although family members provide a low percentage of child care in South Korea, such help is likely to be a useful resource for second childbirth among employed women. Somewhat expectedly, older age and longer time since first birth had negative effects on women's second-child intention. There is growing evidence that, in the lowest fertility societies, paternal investment may be an essential resource for promoting future reproductive behavior of women, especially employed women.     

Maternal investment in a spider with suicidal maternal care, Stegodyphus lineatus (Araneae, Eresidae)

Providing parental care is costly for the parent, but generally beneficial for the young whose survival, growth and reproductive value can be increased. Selection should strongly favour an optimal distribution of parental resources, depending on the relationship between the costs and benefits for parents and their offspring. Parental care is characterized by trade offs in investment, for example between egg size and number of young or providing resources at the egg stage versus the post-hatching stage. Females of the spider Stegodyphus lineatus (Eresidae) produce a single small brood with small eggs and provide the young with regurgitated fluid and later, with their body contents via matriphagy. We asked whether females adjust the investment of resources differentially into eggs, regurgitation feeding and matriphagy, and how maternal investment affects the size of the young at dispersal. We followed the growth of young of broods in the lab and in the field and manipulated brood size in order to determine the pattern of resource allocation. We found that brood size was positively correlated with body mass: larger females had larger broods. Females provided 95% of their body mass to the young, allocating more resources to regurgitation than to matriphagy. Females provided regurgitated food to the young according to the brood size, providing less food when the brood was reduced. Maternal resources had a large influence on offspring mass at dispersal, which is likely to affect their future fitness. The study shows the importance of the female's body mass and her resource allocation decisions for her reproductive outcome.     

An evolutionary perspective on the patterning of maternal investment in pregnancy

Pregnancy is thought to be a metabolically very expensive endeavor, yet investigations have produced inconsistent results concerning the responsiveness of human birth weight to maternal nutritional stress or nutritional intervention. These findings have led some researchers to conclude that fetal growth is strongly buffered against fluctuations in maternal energy balance, making the fetus in effect a “nearly perfect parasite.” This buffering would appear to be a reasonable adaptive response given the high risk of morbidity and mortality associated with low birth weight. However, a life-history approach leads to the prediction that maternal investment strategies in pregnancy should be geared toward maximizing lifetime reproductive success rather than simply the success of the current pregnancy, and by extension that maternal investment strategies should vary with reproductive value. The physiology of human pregnancy in fact appears to include a number of mechanisms that protect maternal energy resources from diversion to the fetus and preserve them for future reproductive events. These mechanisms include adjustment of blood flow to the uterus and perhaps minor adjustments in gestation length, although evidence for the latter is scant. Suggestions are made for ways of investigating these maternal options.     

Different maternal investment strategies for male and female calves in a polygynous mammal

Mater nal effects occur whe n the phe no type of the mother in flue nces that of the young to the detriment of her survival, growth or fitn ess. The in vestment of the mother can be affected by mater nal body condition and/or experience. Trivers-Willard Hypothesis (TWH) and Local Resource Competition Hypothesis (LRCH) are the main hypotheses used to explain bias in birth sex-ratios in mammals, as well as for sex-biased maternal investment. Both hypotheses suggest that a different amount of investment must be expected according to the sex of the young. However, recent studies suggest that these differences are not in quantity but in the strategies: mechanisms and objectives may differ for each sex. We studied how maternal characteristics (age, body mass, body condition, and domi nance status) influence re leva nt aspects of the birth and early growth of the calf (birth date, birth body mass, body mass at weaning, and body condition at weaning) separately for each sex;and how that investment is mediated by milk production and composition (lactose, fat, and protein). One hundred eighty-eight newborns from 75 captive red deer hinds aged from 2 to 19 years were analyzed. The main differential investment observed was related to birth date: when producing a female, hinds give birth earlier in the season only if they have a good body condition;however, when gestating a male it is the older hinds those which deliver earlier. Subsequently, milk production and composition are correlated with birth body mass in female calves, but to weaning body mass in males. Thus, only hind body mass affects the weaning body mass of female calves, compared with age and hind body mass in males. These results suggest that while TWH fits the maternal investment strategy found for male calves, it is LRCH which correlates with the maternal investment patterns observed for females.     

Contraceptive practices and fertility among Southeast Asian,black, and white mothers attending a maternal infant care program

Abstract

The purpose of this analysis was to determine if there were differences in selected fertility characteristics including parity, pregnancy spacing, age at first pregnancy, age of menarche, breastfeeding postpartum, and contraceptive practices among white, black, Hmong, and other Southeast Asian mothers attending a maternal infant care program in Minneapolis, Minnesota, during 1980–82.

White and black mothers were younger than the Hmong and other Asian mothers. The lowest mean age of first pregnancy was among blacks. Ages of first pregnancy were similar for whites, Hmong, and other Asians, although the mean age of menarche was approximately two years later for Hmong and other Southeast Asian mothers compared to the white and black mothers.

Based on self reports at the first postpartum visit one month after delivery, 39 per cent of the whites and 25 per cent of the blacks were breastfeeding. In comparison 8.7 per cent of the Hmong and 17 per cent of the other southeast Asian mothers were breastfeeding.

Maternal age and age at first pregnancy were significant predictors of parity for whites, blacks, Hmong, and the other Southeast Asians. Ever‐use of contraception was a significant predictor of parity only for Hmong.

The highest proportion of ever‐users of contraception was among the white mothers (80 per cent) followed by the blacks (69.3 per cent) and other Asian mothers (34.8 per cent). Hmong mothers had the lowest proportion having used contraception (17.1 per cent).     

Contraceptive practices and fertility among Southeast Asian, black, and white mothers attending a maternal infant care program

The purpose of this analysis was to determine if there were differences in selected fertility characteristics including parity, pregnancy spacing, age at 1st pregnancy, age of menarche, breastfeeding postpartum, and contraceptive practices among white, black, Hmong, and other Southeast Asian mothers attending a maternal infant care program in Minneapolis, Minnesota, during 1980-1982. White and black mothers were younger than the Hmong and other Asian mothers. The lowest mean age of 1st pregnancy was among blacks. Ages of 1st pregnancy were similar for whites, Hmong, and other Asians, although the mean age of menarche was approximately 2 years later for Hmong and other Southeast Asian mothers compared to the white and black mothers. Based on self reports at the 1st postpartum visit 1 month after delivery, 39% of the whites and 25% of the blacks were breastfeeding. In comparison 8.7% of the Hmong and 17% of the other southeast Asian mothers were breastfeeding. Maternal aged age at 1st pregnancy were significant predictors of parity for whites, blacks, Hmong, and other Southeast Asians. Ever-use of contraception was significant predictor of parity only for Hmong. The highest proportion of ever-users of contraception was among the white mothers (80%) followed by the blacks (69.3%) and other Asian mothers (34.85). Hmong mothers had the lowest proportion having used contraception (17.1%).