Lifetime reproductive effort in humans

Lifetime reproductive effort (LRE) measures the total amount of metabolized energy diverted to reproduction during the lifespan. LRE captures key components of the life history and is particularly useful for describing and comparing the life histories of different organisms. Given a simple energetic production constraint, LRE is predicted to be similar in value for very different life histories. However, humans have some unique ecological characteristics that may alter LRE, such as the long post-reproductive lifespan, lengthy juvenile period and the cooperative nature of human foraging and reproduction. We calculate LRE for natural fertility human populations, compare the findings to other mammals and discuss the implications for human life-history evolution. We find that human life-history traits combine to yield the theoretically predicted value (approx. 1.4). Thus, even with the subsidized energy budget and uniqueness of the adult lifespan, human reproductive strategies converge on the same optimal value of LRE. This suggests that the fundamental demographic variables contained in LRE trade-off against one another in a predictable and highly constrained manner.     

Terminal reproductive effort in a marsupial

Life-history theory predicts that as organisms approach the end of their life, they should increase their reproductive effort (RE). However, studies on mammals often find that measures of RE do not vary with maternal age. This might be because offspring have some control over energy transfer which may constrain adaptive variation in RE by mothers, particularly in eutherian mammals where placental function is primarily controlled by offspring. However, in marsupials, energy transfer is primarily by lactation and under maternal control, leaving marsupial mothers free to vary RE. Here, we provide the first analysis, to our knowledge, of age-specific RE in a marsupial, the common brushtail possum. RE, measured as the proportion of maternal mass lost during lactation, was strongly correlated with offspring mass as a yearling. Older females had higher RE, gave birth earlier in the season and were more likely to produce two offspring in a year. Females with high RE in one year were lighter at the beginning of the next breeding season. These results provide the clearest support yet for terminal RE in a mammal.     

Lifetime reproductive success of female mountain gorillas

Studies of lifetime reproductive success (LRS) are important for understanding population dynamics and life history strategies, yet relatively little information is available for long-lived species. This study provides a preliminary assessment of LRS among female mountain gorillas in the Virunga volcanoes region. Adult females produced an average of 3.6 ± 2.1 surviving offspring during their lifetime, which indicates a growing population that contrasts with most other great apes. The standardized variance in LRS (variance/mean(2) = 0.34) was lower than many other mammals and birds. When we excluded the most apparent source of environmental variability (poaching), the average LRS increased to 4.3 ± 1.8 and the standardized variance dropped in half. Adult lifespan was a greater source of variance in LRS than fertility or offspring survival. Females with higher LRS had significantly longer adult lifespans and higher dominance ranks. Results for LRS were similar to another standard fitness measurement, the individually estimated finite rate of increase (λ(ind) ), but λ(ind) showed diminishing benefits for greater longevity.     

Lifetime reproductive success in female Japanese macaques

Lifetime reproductive success, measured by the number of offspring surviving to age five, varied from 0 to 10 in a group of 33 provisioned female Japanese macaques. Of the three contributors to reproductive success, the number of reproductive years, fecundity per year and survivorship of offspring to reproductive age, the first accounted for two-thirds of the variation. Fecundity per year and survivorship were negatively correlated, indicating reproductive costs of reducing interbirth interval. No other demographic measure used, nor the behavioral measure 'dominance rank', significantly correlated with lifetime reproductive success or its components. Age-specific changes in fecundity and infant survival were not found for this sample, neither could cessation of reproduction, even in very old females, be demonstrated.     

Pollen limitation of reproductive effort in willows

Summary Pollen limitation of seed set differs from resource limitation in its implications for the evolution of floral traits. Willow flowers attract insects, but also abundantly produce wind-dispersed pollen. I demonstrated pollen limitation in single branches bearing 2–4 inflorescences (catkins) in a field experiment with five species by artificially increasing or decreasing the pollen load. Because the responses by single branches might be explained by diversion of resources to better-pollinated branches within a plant, a second experiment with one species tested both pollen limitation of whole plants and the autonomy of catkins. Seed set of single willow catkins is unaffected by experimental alterations of seed set in other catkins on the same plant. Hand-pollination of single catkins and of whole plants increased seed set to the same degree, suggesting there is little or no competition for resources between catkins only 5–10 cm apart. Thus, seed set in willows appears to be pollen limited, favoring insect pollination and the evolution of entomophilous traits. The data support previous views that willows have a dual pollination system utilizing wind and insects.     

Respiration and reproductive effort in Xanthium canadense

* BACKGROUND AND AIMS: The proportion of resources devoted to reproduction in the plant is called the reproductive effort (RE), which is most commonly expressed as the proportion of reproductive biomass to total plant biomass production (RE(W)). Reproductive yield is the outcome of photosynthates allocated to reproductive structures minus subsequent respiratory consumption for construction and maintenance of reproductive structures. Thus, RE(W) can differ from RE in terms of photosynthates allocated to reproductive structures (RE(P)). * METHODS: Dry mass growth and respiration of vegetative and reproductive organs were measured in Xanthium canadense and the amount of photosynthates and its partitioning to dry mass growth and respiratory consumption were determined. Differences between RE(W) and RE(P) were analysed in terms of growth and maintenance respiration. * KEY RESULTS: The fraction of allocated photosynthates that was consumed by respiration was smaller in the reproductive organ than in the vegetative organs. Consequently, RE(P) was smaller than RE(W). The smaller respiratory consumption in the reproductive organ resulted from its shorter period of existence and a seasonal decline in temperature, as well as a slower rate of maintenance respiration, although the fraction of photosynthates consumed by growth respiration was larger than in the vegetative organs. * CONCLUSIONS: Reproductive effort in terms of photosynthates (RE(P)) was smaller than that in terms of biomass (RE(W)). This difference resulted from respiratory consumption for maintenance, which was far smaller in the reproductive organ than in vegetative organs.     

Testosterone and reproductive effort in male primates

Considerable evidence suggests that the steroid hormone testosterone mediates major life-history trade-offs in vertebrates, promoting mating effort at the expense of parenting effort or survival. Observations from a range of wild primates support the “Challenge Hypothesis,” which posits that variation in male testosterone is more closely associated with aggressive mating competition than with reproductive physiology. In both seasonally and non-seasonally breeding species, males increase testosterone production primarily when competing for fecund females. In species where males compete to maintain long-term access to females, testosterone increases when males are threatened with losing access to females, rather than during mating periods. And when male status is linked to mating success, and dependent on aggression, high-ranking males normally maintain higher testosterone levels than subordinates, particularly when dominance hierarchies are unstable. Trade-offs between parenting effort and mating effort appear to be weak in most primates, because direct investment in the form of infant transport and provisioning is rare. Instead, infant protection is the primary form of paternal investment in the order. Testosterone does not inhibit this form of investment, which relies on male aggression. Testosterone has a wide range of effects in primates that plausibly function to support male competitive behavior. These include psychological effects related to dominance striving, analgesic effects, and effects on the development and maintenance of the armaments and adornments that males employ in mating competition.     

Lifetime reproductive success in reproductively suppressed female voles

A population of the grey red-backed vole, Clethrionomys rufocanus bedfordiae, was investigated on a 1 ha control grid and a 1 ha grid on which the voles were fed within a 2.1 ha outdoor enclosure in Hokkaido, Japan by live trapping from 1984 to 1986, for testing the Reproductive Suppression Model of Wasser and Barash (1983)-females can optimize their lifetime reproductive success by suppressing reproduction when future conditions for the survival of offspring are likely to be sufficiently better than present ones as to exceed the costs of the suppression itself. Age at the first pregnancy more varied in a higher density population on the experimental grid and females could be classified into the early and the late reproductive type in two generations (A: females born from February to June 1985; B: females born from September to November 1985). Lifetime reproductive success (the number of pregnancies, the number of successful litters, and the number of offspring) was not different between the early and the late reproducing females. The late reproducing females lived for longer periods than the early reproducing females, so that the loss by delayed start of reproduction was compensated for by a longer life span. Life span was not different between offspring of the early and the late reproducing females. These facts supported the Reproductive Suppression Model.     

Genetic variability and reproductive effort in Polygonum pensylvanicum

Six populations of the annual plant Polygonum pensylvanicum were analysed for isozyme variability and reproductive effort. The populations were found to be similar in both their isozyme constituents and in their mean reproductive efforts. Mean reproductive effort did not correlate with mean population heterozygosity nor did individual reproductive effort correlate with individual heterozygosity. Polygonum pensylvanicum exhibits several of the features which have been predicted for weedy plant species.     

Modelling reproductive effort in sub-and maritime Antarctic mosses

A comparison is made of the reproductive effort (RE), considered as the investment in sporophyte relative to gametophyte biomass, of eight species of moss occurring at sub-and maritime Antarctic sites. Six of the species showed smaller sporophytes and game-tophytes at the climatically more extreme maritime Antaretic sites and one species showed no size difference between regions. The remaining species, although showing no regional difference, showed some evidence of a reverse pattern, with higher altitude samples having greater biomass than lower altitude samples. Spore counts indicated a measure of compensation in maritime Antarctic samples, with no significant decrease in spore output in several species despite smaller sporophyte biomass. The relationship between sporophyte (S) and gametophyte (G) biomass within samples was described by an allometric curve (S=aG ^b ) which gave a better fit than a straight line for six species. This form of model allows comparisons of patterns of RE to be made between samples with non-or partially overlapping size distributions, even when the relationship involves size-dependence. An allometric curve was not appropriate for describing samples of one species (Andreaea regularis), and insufficient data were available to identify any relationship in Polytrichum alpinum. The exponent (b) differed between species, but there were no statistically significant differences between exponents from samples of the same species. Samples of two species could further be described by the same coefficient (a), indicating that they lie on the same curve. However, samples of three species from sub-Antarctic South Georgia gave significantly higher coefficients, indicating increased RE relative to maritime Antarctic populations.     

Lifetime reproductive success and density-dependent, multi-variable resource selection

Individuals are predicted to maximize lifetime reproductive success (LRS) through selective use of resources; however, a wide range of ecological and social processes may prevent individuals from always using the highest-quality resources available. Resource selection functions (RSFs) estimate the relative amount of time an individual spends using a resource as a function of the proportional availability of that resource. We quantified the association between LRS and coefficients of individual-based RSFs describing lifetime resource selection for 267 female red deer (Cervus elaphus) of the Isle of Rum, Scotland, from 1970 to 2001. LRS was significantly related to first- and second-order effects of selection for Agrostis/Festuca grassland and proximity to the sea coast (quality of forage within Agrostis/Festuca grassland was highest nearest the coast (ratio of short:long grassland)). The benefits of selecting for quality in Agrostis/Festuca grassland, however, traded-off with increases in LRS gained by avoiding conspecific density. LRS was inversely associated with local density, which was highest along the coast, and reproductive benefits of selecting Agrostis/Festuca grassland diminished with increasing density. We discuss the relevance of these results to our understanding of the spatial distribution of red deer abundance, and potential applications of our approach to evolutionary and applied ecology.     

Dietary phosphorus availability influences female cricket lifetime reproductive effort

1. Recent ecological stoichiometric findings indicate that the relationships among key macronutrient elements [e.g. carbon (C), nitrogen (N), and phosphorus (P) of organisms and their resources] may underlie variation in life‐history traits. The amount of phosphorus in an individual's body is often correlated with its rate of growth, and low‐phosphorus diets are known to reduce growth in a number of insect and crustacean herbivores. 2. These findings suggest that the stoichiometric imbalance between organismal biomass requirements and the relative scarcity of nutrients in nature may also underlie variation in lifetime reproductive success. 3. This study investigated how dietary phosphorus availability during adulthood influenced lifetime reproductive effort, compensatory feeding, lifespan, condition, and stoichiometry of adult European House Cricket, Acheta domesticus. 4. Female crickets fed high amounts of phosphorus during adulthood laid significantly more eggs compared to those fed low amounts of phosphorus. Phosphorus availability did not directly influence lifespan, condition, or body stoichiometry, and crickets did not compensate for low phosphorus diets by eating more food. 5. A stoichiometric perspective may help understand the causes of variation in invertebrate fitness.     

Seagrass reproductive effort as an ecological indicator of disturbance

The available information on the changes in the reproductive effort (RE) of seagrasses in response to disturbances was reviewed and analysed to assess if seagrasses invest in RE when disturbed, and if this response is related to specific types of disturbance or seagrass traits. In 72% of the documented cases RE increased with disturbance, in 25% it decreased, and in 3% no changes were reported. Overall, seagrass RE increased 4-fold with disturbance. Anthropogenic disturbances had the highest impact on RE (a 13-fold increase); 3 times higher than the effect of natural disturbances. Mechanical and sedimentary/hydrodynamics disturbances caused the highest RE increase (9- and 5-fold, respectively). The positive RE response was significantly correlated with rhizome diameter of seagrasses, but not with shoot size (mass or length), suggesting that species with higher storage capacity have a higher capacity of investing in sexual reproduction when conditions deteriorate. Seagrasses showed a general trend of increasing RE under disturbance; this was evident regardless of the origin and type of disturbance, which suggests that changes in seagrass RE provide a valuable indicator of disturbance in coastal areas.     

Age- and density-dependent reproductive effort in male red deer

Reproductive effort in female ungulates originates from gestation and lactation and has been studied extensively; however, no comparable studies of reproductive effort in males (due to fighting for access to mates) have, to our knowledge, previously been reported. Here, we report on weight loss of male red deer during the annual mating season--a direct measure of male reproductive effort (or somatic reproductive costs). The 'terminal investment' hypothesis predicts that reproductive effort should increase with age, given that costs remain stable. We also propose the 'mating strategy-effort' hypothesis, which predicts that reproductive effort peaks in prime-aged males, since they are most often the harem holders. Consistent with the mating strategy-effort hypothesis, relative weight loss during the rutting season peaked at prime age and was lower in younger and senescent males. Weight loss during the rut was relatively smaller as density increased and more so for older males. This is probably primarily due to males (particularly senescent males) starting their rut in poorer condition at high density. The pattern of reproductive effort in males with regard to age and density therefore differs markedly from the pattern reported for females.     

Manipulating reproductive effort leads to changes in female reproductive scheduling but not oxidative stress

The trade‐off between reproductive investment and lifespan is the single most important concept in life‐history theory. A variety of sources of evidence support the existence of this trade‐off, but the physiological costs of reproduction that underlie this relationship remain poorly understood. The Free Radical Theory of Ageing suggests that oxidative stress, which occurs when there is an imbalance between the production of damaging Reactive Oxygen Species (ROS) and protective antioxidants, may be an important mediator of this trade‐off. We sought to test this theory by manipulating the reproductive investment of female mice (Mus musculus domesticus) and measuring the effects on a number of life history and oxidative stress variables. Females with a greater reproductive load showed no consistent increase in oxidative damage above females who had a smaller reproductive load. The groups differed, however, in their food consumption, reproductive scheduling and mean offspring mass. Of particular note, females with a very high reproductive load delayed blastocyst implantation of their second litter, potentially mitigating the costs of energetically costly reproductive periods. Our results highlight that females use strategies to offset particularly costly periods of reproduction and illustrate the absence of a simple relationship between oxidative stress and reproduction.