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Increased infertility with age in men and women.

OBJECTIVE: To estimate the effects of aging on the percentage of outwardly healthy couples who are sterile (completely unable to conceive without assisted reproduction) or infertile (unable to conceive within a year of unprotected intercourse). METHODS: A prospective fecundability study was conducted in a sample of 782 couples recruited from 7 European centers for natural family planning. Women aged 18-40 years were eligible. Daily intercourse records were used to adjust for timing and frequency of intercourse when estimating the per-menstrual-cycle probability of conception. The number of menstrual cycles required to conceive a clinical pregnancy and the probability of sterility and infertility were derived from the estimated fecundability distributions for men and women of different ages. RESULTS: Sterility was estimated at about 1%; this percent did not change with age. The percentage infertility was estimated at 8% for women aged 19-26 years, 13-14% for women aged 27-34 years and 18% for women aged 35-39 years. Starting in the late 30s, male age was an important factor, with the percentage failing to conceive within 12 cycles increasing from an estimated 18-28% between ages 35 and 40 years. The estimated percentage of infertile couples that would be able to conceive after an additional 12 cycles of trying varied from 43-63% depending on age. CONCLUSION: Increased infertility in older couples is attributable primarily to declines in fertility rates rather than to absolute sterility. Many infertile couples will conceive if they try for an additional year.

Adolescent↗

Reproductive effects of male psychologic stress.

OBJECTIVES: Male psychologic stress can affect semen quality and couple fecundability. METHODS: We conducted a prospective study of 430 Danish couples who were trying to become pregnant for the first time. In each menstrual cycle, both partners filled out a 12-item version of the General Health Questionnaire (GHQ-12), an assessment of psychologic stress. A blood sample was drawn from both partners. The men collected a semen sample at enrollment and each month during follow up. RESULTS: Semen quality was not related to the man's GHQ score in either within-subject analyses or between-subject analyses. There were no consistent associations between stress and serum concentration of luteinizing hormone, follicle-stimulating hormone, inhibin B, testosterone, or estradiol. The pregnancy rate was 14% in cycles with an absolute GHQ-12 score in the highest quartile, compared with 18% in low-stress cycles. Odds of pregnancy decreased moderately with increasing score. The effect was confined to 77 men with a sperm density below 20 million/mL (adjusted odds ratio = 0.06; 95% confidence interval = 1.01-0.58 for highest distressed quartile vs. lowest distressed quartile in this low sperm density group). In within-subject analyses, cycle-specific changes in male stress did not change the odds of pregnancy. High absolute score was associated with a lower frequency of sexual intercourse, but adjustment for frequency had little effect. CONCLUSION: The effect of a man's daily life psychologic stress on his semen quality is small or nonexistent. Our results indicate an effect of stress only on fecundability, and this only among men with low sperm concentration.

Adult↗

Has human fertility declined over time?: why we may never know.

BACKGROUND: Reports of decreased semen quality over time have raised concerns about possible reductions in human fertility. Studies of couple fertility have produced conflicting results. We evaluate how changes in the availability and use of effective contraception and induced abortion might bias the direct study of time trends in couple fertility. METHODS: We assess the potential for bias in the context of 2 common study designs: (1) a study of time-to-pregnancy that estimates fecundability (excluding unintended pregnancies) and (2) a study of infertility rates that categorizes couples as fertile or infertile (including couples with unintended pregnancies as fertile). RESULTS: In time-to-pregnancy studies, bias alone could produce more than a 2-fold apparent increase in fecundability over recent decades. In studies of infertility rates, the bias works in the opposite direction: a 30% underestimation of infertility during earlier decades could produce an apparent decrease in fertility over time. CONCLUSIONS: Over the past 5 decades, changes in social factors that affect the rate and fate of unintended pregnancies could substantially bias time trends in fertility. These biases may explain the conflicting reports in the literature. Except in rare settings in which the factors affecting reproductive choices have not changed, it is probably impossible to identify biologic changes in fertility over recent decades.

Abortion, Induced↗

The influence of dietary protein on the experimental epidemiology of Heligmosomoides polygyrus (Nematoda) in the laboratory mouse.

The influence of dietary protein on the epidemiology of an intestinal helminth infection was investigated with an experimental system that allowed transmission of the nematode Heligmosomoides polygyrus to occur naturally between laboratory mice. Mortality of mice was greatly increased in infected populations that were fed ad libitum on synthetic diets containing 2% compared with 16% protein. Larger numbers of larval and adult H. polygyrus were found to infect mice in the low-protein cage compared with the high-protein cage. No evidence for density dependence in the fecundity of female worms was detected; on average the daily egg output per female worm was greater for parasites infecting mice in the low-protein cage. The rate at which naïve mice acquired infection was also higher in the low-protein cage. Pinworm (Aspiculuris tetraptera) became established in each cage, and average worm burdens were again greater in the low-protein cage. The acquisition of resistance to reinfection was not found to be an important factor influencing the survival of parasites infecting mice in either cage. The epidemiology of H. polygyrus and A. tetraptera was therefore characterized by low average worm burdens and high host survival in a well-nourished population of mice, and by a high intensity of infection and severe parasite-induced host mortality in a malnourished colony of mice. This reflects differences in the survival and fecundity of adult parasites between mice in the two cages, and suggests that malnourished mice are predisposed to acquire large numbers of several species of intestinal worm.

Animals↗

Diagnosing senescence: inferring evolutionary causes from phenotypic patterns can be misleading.

Based on the predictions of two theories for the evolution of senescence, the 'antagonistic pleiotropy' and the 'mutation accumulation' theory, an age-specific increase in mortality and a decrease in fecundity are widely used criteria to diagnose senescence in natural, and laboratory populations. In this study we question the reliability of these criteria. Using a simple model we show that similar phenotypic patterns result from optimal life histories without senescence. With a tradeoff between reproduction and period survival, optimal life histories produce patterns of increasing mortality and decreasing fecundity as organisms age, even if the tradeoff does not deteriorate with age, so that we are not forced to invoke genetic effects such as antagonistic pleiotropy or accumulation of deleterious mutations to explain such patterns. Furthermore, if optimal life history theory is applied to senescent organisms, phenotypic patterns can result that are usually not associated with senescence. We conclude that the reliability of a diagnosis of senescence based on phenotypic patterns and the comprehension of the phenomenon senescence depends critically on understanding to what extent tradeoffs are determined by the effects of segregating genes.

Aging↗

Host-parasite coevolution: comparative evidence for covariation of life history traits in primates and oxyurid parasites.

The environmental factors that drive the evolution of parasite life histories are mostly unknown. Given that hosts provide the principal environmental features parasites have to deal with, and given that these features (such as resource availability and immune responses) are well characterized by the life history of the host, we may expect natural selection to result in covariation between parasite and host life histories. Moreover, some parasites show a high degree of host specificity, and cladistic analyses have shown that host and parasite phylogenies can be highly congruent. These considerations suggest that parasite and host life histories may covary. The central argument in the theory of life history evolution concerns the existence of trade-offs between traits. For parasitic nematodes it has been shown that larger body sizes induce higher fecundity, but this is achieved at the expense of delayed maturity. As high adult mortality would select for reduced age at maturity, the selective benefit of increased fecundity is expressed only if adult mortality is low. Parasite adult mortality may depend on a number of factors, including host longevity. Here we tested the hypothesis concerning the positive covariation between parasite body size (which reflects parasite longevity) and host longevity. To achieve this goal, we used the association between the pinworms (Oxyuridae, Nematoda) and their primate hosts. Oxyurids are highly host specific and are supposed to be involved in a coevolutionary process with their hosts. We found that female parasite body length was positively correlated with host longevity after correcting for phylogeny and host body mass. Conversely, male parasite body length and host longevity were not correlated. These results confirm that host longevity may represent a constraint on the evolution of body size in oxyurids, at least in females. The discrepancy between female and male oxyurids is likely to depend on the particular mode of reproduction of this taxon (haplodiploidy), which should result in weak (or even null) selection pressures to an increase of body size in males.

Animals↗

Postponed reproduction as an adaptation to winter conditions in Drosophila melanogaster: evidence for clinal variation under semi-natural conditions.

Patterns of climatic adaptation in drosophila and other insects have largely been inferred from laboratory comparisons of traits that vary clinally. Here, we extend this research to comparisons under semi-natural conditions. To test for clinal variation in reproductive patterns and survival over winter, Drosophila melanogaster populations were initiated from seven collection sites along the eastern coast of Australia, ranging from tropical to temperate regions. The fecundity and survival of these populations were monitored in field cages at a temperate location until all adults had died more than 5 months later. Total fecundity showed a curvilinear relationship with latitude, due to higher egg production by high- and low-latitude populations. Adults from temperate locations survived winter conditions better than those from subtropical populations but not tropical ones. There was a linear cline in the timing of egg production: temperate populations produced eggs later than populations from lower latitudes. This cline is likely to be adaptive because egg-to-adult viability experiments indicated that only eggs laid in spring developed successfully to the adult stage. There was no evidence for climatic adaptation in the immature stages. The adult mortality rate increased gradually over winter, and in some populations was also correlated with the minimum ambient temperature. These results indicate that adaptation to winter conditions in D. melanogaster has involved shifts in reproductive patterns.

Acclimatization↗

The role of parasites in the dynamics of a reindeer population.

Even though theoretical models show that parasites may regulate host population densities, few empirical studies have given support to this hypothesis. We present experimental and observational evidence for a host-parasite interaction where the parasite has sufficient impact on host population dynamics for regulation to occur. During a six year study of the Svalbard reindeer and its parasitic gastrointestinal nematode Ostertagia gruehneri we found that anthelminthic treatment in April-May increased the probability of a reindeer having a calf in the next year, compared with untreated controls. However, treatment did not influence the over-winter survival of the reindeer. The annual variation in the degree to which parasites depressed fecundity was positively related to the abundance of O. gruehneri infection the previous October, which in turn was related to host density two years earlier. In addition to the treatment effect, there was a strong negative effect of winter precipitation on the probability of female reindeer having a calf. A simple matrix model was parameterized using estimates from our experimental and observational data. This model shows that the parasite-mediated effect on fecundity was sufficient to regulate reindeer densities around observed host densities.

Animal Diseases↗

Harm to females increases with male body size in Drosophila melanogaster.

Previous studies indicate that female Drosophila melanogaster are harmed by their mates through copulation. Here, we demonstrate that the harm that males inflict upon females increases with male size. Specifically, both the lifespan and egg-production rate of females decreased significantly as an increasing function of the body size of their mates. Consequently, females mating with larger males had lower lifetime fitness. The detrimental effect of male size on female longevity was not mediated by male effects on female fecundity, egg-production rate or female-remating behaviour. Similarly, the influence of male size on female lifetime fecundity was independent of the male-size effect on female longevity. There was no relationship between female size and female resistance to male harm. Thus, although increasing male body size is known to enhance male mating success, it has a detrimental effect on the direct fitness of their mates. Our results indicate that this harm is a pleiotropic effect of some other selected function and not an adaptation. To the extent that females prefer to mate with larger males, this choice is harmful, a pattern that is consistent with the theory of sexually antagonistic coevolution.

Animals↗

Life-history correlates of maximum population growth rates in marine fishes.

Theory predicts that populations of animals with late maturity, low fecundity, large body size and low body growth rates will have low potential rates of population increase at low abundance. If this is true, then these traits may be used to predict the intrinsic rate of increase for species or populations, as well as extinction risks. We used life-history and population data for 63 stocks of commercially exploited fish species from the northeast Atlantic to test relationships between life-history parameters and the rate of population increase at low abundance. We used cross-taxonomic analyses among stocks and among species, and analyses that accounted for phylogenetic relationships. These analyses confirmed that large-bodied, slow-growing stocks and species had significantly lower rates of recruitment and adult production per spawning adult at low abundance. Furthermore, high ages at maturity were significantly correlated with low maximum recruit production. Contrary to expectation, fecundity was significantly negatively related to recruit production, due to its positive relationship with maximum body size. Our results support theoretical predictions, and suggest that a simply measured life-history parameter can provide a useful tool for predicting rates of recovery from low population abundance.

Animals↗

Copulatory courtship signals male genetic quality in cucumber beetles.

In the spotted cucumber beetle, Diabrotica undecimpunctata howardi (Coleoptera: Chrysomelidae), males court females during copulation by stroking them with their antennae. Stroking occurs exclusively during the first stages of copulation, after a male has penetrated a female's vaginal duct but before he is allowed access to her bursa copulatrix. Females accept the spermatophore of fast-stroking males and reject those of slow-stroking males by relaxing or constricting muscles distorting the vaginal duct. Here, we measure the repeatability of stroking behaviour within males, examine the effect of losing one antenna on male attractiveness and test whether such female control results in direct phenotypic benefits for the discriminating female or indirect genetic benefits that appear in her offspring. We also use a half-sibling design to quantify the variance and heritability of stroking speed and endurance. Female beetles were paired with a male that was known to stroke either quickly or slowly. No difference was found in the resulting fecundity or egg-hatching rate of the females, or in the survivorship, development rate, size, age at first reproduction or fecundity of their offspring indicating that no direct benefits are gained by discriminating among males on the basis of stroking speed. There were, however, good-genes benefits for the mates of fast-stroking males. Offspring of fast-stroking fathers were also fast strokers and were more likely to be accepted as mates than offspring of slow-stroking fathers. There was substantial variance among sires in stroking speed and endurance and the heritability of each trait was high. The antennal stroking rate was highly repeatable in successive mating attempts and males with only one antenna were not accepted as mates. The repeatability within males, variability between males and heritability between generations of copulatory stroking combine to provide females with a reliable and honest signal of the genetic quality of courting males.

Animals↗

Functional consequences of genetic diversity in Strongyloides ratti infections.

Parasitic nematodes show levels of genetic diversity comparable to other taxa, but the functional consequences of this are not understood. Thus, a large body of theoretical work highlights the potential consequences of parasite genetic diversity for the epidemiology of parasite infections and its possible implications for the evolution of host and parasite populations. However, few relevant empirical data are available from parasites in general and none from parasitic nematodes in particular. Here, we test two hypotheses. First, that different parasitic nematode genotypes vary in life-history traits, such as survivorship and fecundity, which may cause variation in infection dynamics. Second, that different parasitic nematode genotypes interact within the host (either directly or via the host immune system) to increase the mean reproductive output of mixed-genotype infections compared with single-genotype infections. We test these hypotheses in laboratory infections using genetically homogeneous lines of Strongyloides ratti. We find that nematode genotypes do vary in their survivorship and fecundity and, consequently, in their dynamics of infection. However, we find little evidence of interactions between genotypes within hosts under a variety of trickle- and single-infected infection regimes.

Animals↗

Life-history strategies associated with local population variability confer regional stability.

A widely held ecological tenet is that, at the local scale, populations of K-selected species (i.e. low fecundity, long lifespan and large body size) will be less variable than populations of r-selected species (i.e. high fecundity, short lifespan and small body size). We examined the relationship between long-term population trends and life-history attributes for 185 bird species in the Czech Republic and found that, at regional spatial scales and over moderate temporal scales (100-120 years), K-selected bird species were more likely to show both large increases and decreases in population size than r-selected species. We conclude that life-history attributes commonly associated with variable populations at the local scale, confer stability at the regional scale.

Animals↗

The energetic budget of Anopheles stephensi infected with Plasmodium chabaudi: is energy depletion a mechanism for virulence?

Evidence continues to accumulate showing that the malaria parasites (Plasmodium spp.) reduce the survival and fecundity of their mosquito vectors (Anopheles spp.). Our ability to identify the possible epidemiological and evolutionary consequences of these parasite-induced fitness reductions has been hampered by a poor understanding of the physiological basis of these shifts. Here, we explore whether the reductions in fecundity and longevity are the result of a parasite-mediated depletion or reallocation of the energetic resources of the mosquito. Mosquitoes infected with Plasmodium chabaudi were expected to have less energetic resources than uninfected mosquitoes, and energy levels were predicted to be lowest in mosquitoes infected with the most virulent parasite genotypes. Not only was there no evidence of a parasite-mediated reduction in the overall energetic budget of mosquitoes, but Plasmodium was actually associated with increased levels of glucose, a key insect nutritional and energetic resource. The data strongly suggest the existence of an increase in sugar feeding in mosquitoes infected with Plasmodium. We suggest different adaptive explanations for an enhanced sugar uptake in infected mosquitoes and call for more studies to investigate the physiological role of glucose in the Plasmodium-mosquito interaction.

Adaptation, Physiological↗

Paternal investment directly affects female reproductive effort in an insect.

Female reproductive effort can be influenced by the quality of her mate. In some species, females increase their reproductive effort by differentially allocating resources after mating with high-quality males. Examination of female reproductive effort in relation to male quality has implications for estimating the evolvability of traits and for sexual-selection models. Accurate quantification of reproductive investment is not possible in many species. Butterflies are an exception, as most nectar-feeding species emerge with almost intact reproductive resources, and in some species males provide nutrients at mating that enhance female fecundity. By manipulating male donations and using radioactive isotopes, we quantified the effect of variation in nutrient provisioning on female reproductive effort in two butterfly species. In the greenveined white butterfly, Pieris napi, females increased their reproductive effort after receiving large male donations. By contrast, in the speckled wood, Pararge aegeria, where males do not provide nutrients, female reproductive effort was independent of male ejaculate. Increased reproductive effort in Pieris napi resulted from the production of more eggs, rather than from investing more resources per egg. In this species donating ability is heritable; hence females laying more eggs after mating with high-donating males benefit both through higher fecundity and through the production of high-donating sons.

Animals↗

Nonlinear and correlational sexual selection on 'honest' female ornamentation.

Female ornamentation has long been overlooked because of the greater prevalence of elaborate displays in males. However, the circumstances under which females would benefit from honestly signalling their quality are limited. Females are not expected to invest in ornamentation unless the fitness benefits of the ornament exceed those derived from investing the resources directly into offspring. It has been proposed that when females gain direct benefits from mating, females may instead be selected for ornamentation that deceives males about their reproductive state. In the empidid dance flies, males frequently provide nuptial gifts and it is usually only the female that is ornamented. Female traits in empidids, such as abdominal sacs and enlarged pinnate leg scales, have been proposed to 'deceive' males into matings by disguising egg maturity. We quantified sexual selection in the dance fly Rhamphomyia tarsata and found escalating, quadratic selection on pinnate scales and that pinnate scales honestly reflect female fecundity. Mated females had a larger total number and more mature eggs than unmated females, highlighting a potential benefit rather than a cost of male mate choice. We also show correlational selection on female pinnate scales and fecundity. Correlational selection, equivalent investment patterns or increased nutrition from nuptial gifts may all maintain honesty in female ornamentation.

Animal Communication↗

Pollen feeding in the butterfly Heliconius charitonia: isotopic evidence for essential amino acid transfer from pollen to eggs.

Heliconius and Laparus butterflies exhibit a unique pollen-collecting behaviour that enhances lifespan and fecundity. The specific nutritional contribution of pollen, however, had not been previously demonstrated. We used stable isotope variation to trace the carbon flow into eggs from corn pollen provided experimentally to ovipositing female Heliconius charitonia, and to evaluate the use of isotopically contrasting nectar sugars in egg amino acids. The delta(13)C of individual amino acids from pollen, larval host plant and the eggs from experimental butterflies was measured with gas chromatography/combustion/isotope ratio mass spectrometry (GC/C/IRMS), to evaluate amino acid transfer. The delta(13)C of egg essential amino acids indicated a transfer of essential amino acids from pollen to butterfly eggs. However, the delta(13)C of non-essential amino acids reflected the isotopic composition of the artificial nectar, indicating that H. charitonia synthesizes non-essential amino acids from dietary sugars. This, to our knowledge, is the first direct demonstration of amino acid transfer from pollen to butterfly eggs, and suggests that essential amino acids in particular are a key resource for extending lifespan and fecundity in Heliconius butterflies.

Amino Acids, Essential↗

Large breasts and narrow waists indicate high reproductive potential in women.

Physical characteristics, such as breast size and waist-to-hip ratio (WHR), function as important features used by human males to assess female attractiveness. Males supposedly pay attention to these features because they serve as cues to fecundity and health. Here, we document that women with higher breast-to-underbreast ratio (large breasts) and women with relatively low WHR (narrow waists) have higher fecundity as assessed by precise measurements of daily levels of 17-beta-oestradiol (E2) and progesterone. Furthermore, women who are characterized by both narrow waists and large breasts have 26% higher mean E2 and 37% higher mean mid-cycle E2 levels than women from three groups with other combinations of body-shape variables, i.e. low WHR with small breasts and high WHR with either large or small breasts. Such gains in hormone levels among the preferred mates may lead to a substantial rise in the probability of conception, thus providing a significant fitness benefit.

Adult↗