Are reproductive and somatic senescence coupled in humans? Late, but not early, reproduction correlated with longevity in historical Sami women

Evolutionary theory of senescence emphasizes the importance of intense selection on early reproduction owing to the declining force of natural selection with age that constrains lifespan. In humans, recent studies have, however, suggested that late-life mortality might be more closely related to late rather than early reproduction, although the role of late reproduction on fitness remains unclear. We examined the association between early and late reproduction with longevity in historical post-reproductive Sami women. We also estimated the strength of natural selection on early and late reproduction using path analysis, and the effect of reproductive timing on offspring survival to adulthood and maternal risk of dying at childbirth. We found that natural selection favoured both earlier start and later cessation of reproduction, and higher total fecundity. Maternal age at childbirth was not related to offspring or maternal survival. Interestingly, females who produced their last offspring at advanced age also lived longest, while age at first reproduction and total fecundity were unrelated to female longevity. Our results thus suggest that reproductive and somatic senescence may have been coupled in these human populations, and that selection could have favoured late reproduction. We discuss alternative hypotheses for the mechanisms which might have promoted the association between late reproduction and longevity.     

Pleiotropic Effects of methoprene-tolerant(Met), a Gene Involved in Juvenile Hormone Metabolism, on Life History Traits in Drosophila melanogaster

Life history theory assumes that there are alleles with pleiotropic effects on fitness components. Although quantitative genetic data are often consistent with pleiotropy, there are few explicit examples of pleiotropic loci. The Drosophila melanogaster gene Methoprene-tolerant (Met) may be such a locus. The Met gene product, a putative juvenile hormone receptor, facilitates the action of juvenile hormone (JH) and JH analogs; JH affects many life history traits in arthropods. Here we use quantitative complementation to investigate effects of Met mutant and wildtype alleles on female developmental time, onset of reproduction, and fecundity. Whereas the alleles did not differ in their effects on developmental time, we detected allelic variation for the onset of reproduction and for age-specific fecundity. Alleles influenced phenotypic co-variances among traits (developmental time and onset of reproduction; onset of reproduction and both early and late fecundity; early and late fecundity), suggesting that alleles of Met vary in their pleiotropic effects upon life history. Furthermore, the genetic covariance between developmental time and early fecundity attributed to alleles of Met was negative, indicating consistent pleiotropic effects among alleles on these traits. The allelic effects of Met support genetic models where pleiotropy at genes associated with hormone regulation can contribute to the evolution of life history traits.     

Temporal structure and some features of stock dynamics of pink salmon <Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis> (Salmonidae)

Characteristics of early and late migrants of pink salmon Oncorhynchus gorbuscha that spawned in the rivers of the coast of Aniva Bay (southern Sakhalin) and Iturup Island (southern Kuril Islands) are described. Upstream migration of the late form is characterized by the presence of large exemplars, especially males, and increasing proportion of these fishes in the catches. Absolute fecundity of the females of the early and late forms can be similar (Iturup Island) or different (Aniva Bay), but relative fecundity is lower in the females of the late form in both regions. Production of a larger number of eggs by the females of the early form is associated with a larger level of mortality: the abundance of the early form is comparatively low, and substantial annual variation of the abundance is registered. Relative abundance of pink salmon from the temporal groups depends mainly on reproductive conditions during the freshwater part of their life cycle. Analysis of size composition of the fish conducted over many years shows a trend in the ratio between the body lengths of males and females: unfavorable foraging conditions (and slower growth rate) lead to the appearance of females that are larger than males. To take into account a positive correlation between the body length and fecundity of females, this feature is regarded as a compensatory reaction of the population directed to more intensive reproduction at poor foraging conditions.     

Reproductive pattern of three amerindian communities from Costa Rica

The bases of the reproductory models of three indigenous communities from Costa Rica belonging to the same cultural area (Chibcha) but with different social, economical and environmental conditions were studied. In this situation it was observed how the study of the data on fertility and fecundity, analyzed together with other reproductory and social characteristics of women, can precisely determine the reproductory model of the communities to which they belong. A homogeneous fecundity was presented in the studied communities, but one which responded to different reproductory patterns. Starting from the multivariate analysis of the fertility variables, the studied women were divided into two groups. One of the groups presents the highest levels of length of breast feeding period and amenorrhea, together with the near total lack of artificial breast feeding and the early introduction of normal feeding. This model is predominant among the Guaymies. The other one, together with the short period of breast feeding and amenorrhea, shows an early introduction of the bottle and a late one of solid food in the feeding process. It is characteristic of the Huetars. This links a certain reproductory model to an indigenous community and a way of life. The reproductory pattern, a clearly biological model, to an ecological and cultural environment.     

An Old Mom Keeps You Young: Mother’s Age at Last Birth and Offspring Longevity in Nineteenth-Century Utah

This study analyzes the intergenerational effects of late childbearing on offspring’s adult longevity in a population in Utah (United States) that does not display evidence of parity-specific birth control—a so-called natural fertility population. Studies have found that for women who experience late menopause and prolonged reproduction, aging is postponed and longevity is increased. This is believed to indicate female “robustness” and the impact of biological or genetic factors. If indeed there is a genetic component involved, one would expect to also find evidence for the intergenerational transmission of longevity benefits. Our study investigates the relationship between prolonged natural fertility of mothers and their offspring’s survival rates in adulthood. Gompertz regression models (N = 7,716) revealed that the offspring of mothers who were naturally fertile until a relatively advanced age lived significantly longer. This observed positive effect of late reproduction was not independent of but conditional upon survival of the mother to the end of her fecundity (defined as age 50). Offspring’s relative risks at death beyond age 50 were 6–12 percent lower than those of their counterparts born to mothers who had an average age at last birth. Our results, which account for various early, adult, and later-life conditions, as well as shared frailty, suggest that there is a positive relationship between mother’s age at last birth and offspring longevity, and strengthen the notion that age at menopause is a good predictor of this relationship.     

Secular Trends in Menarcheal Age in India-Evidence from the Indian Human Development Survey

Background

Evidence from a number of countries in Europe and North America point towards the secular declining trend in menarcheal age with considerable spatial variations over the past two centuries. Similar trends were reported in several developing countries from Asia, Africa and Latin America. However, data corroborating any secular trend in the menarcheal age of the Indian population remained sparse and inadequately verified.

Methods

We examined secular trends, regional heterogeneity and association of socioeconomic, anthropometric and contextual factors with menarcheal age among ever-married women (15–49 years) in India. Using the pseudo cohort data approach, we fit multiple linear regression models to estimate secular trends in menarcheal age of 91394 ever-married women using the Indian Human Development Survey.

Results

The mean age at menarche among Indian women was 13.76 years (95 % CI: 13.75, 13.77) in 2005. It declined by three months from 13.83 years (95% CI: 13.81, 13.85) among women born prior to 1955–1964, to nearly 13.62 years (95% CI: 13.58, 13.67) among women born during late 1985–1989. However, these aggregate national figures mask extensive spatial heterogeneity as mean age at menarche varied from 15.0 years in Himachal Pradesh during 1955–1964 (95% CI: 14.89–15.11) to about 12.1 years in Assam (95% CI: 11.63–12.56) during 1985–1989.

Conclusion

The regression analysis established a reduction of nearly one month per decade, suggesting a secular decline in age at menarche among Indian women. Notably, the menarcheal age was significantly associated with the area of residence, geographic region, linguistic groups, educational attainment, wealth status, caste and religious affiliations among Indian women.     

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

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

Trends and differentials in menarcheal age in China

This study examines trends in menarcheal age of women born in China between 1950 and 1973, and explores the impact of relevant social background characteristics on the timing of first menarche. Data on recalled ages of menarche collected in the 1988 Chinese Two-per-Thousand Fertility Survey are used in a linear regression model where the covariates are transformed with the help of an Alternating Conditional Expectation (ACE) algorithm. The results indicate that a trend towards early menarche has evolved in China during recent decades. The pattern of early menarche is especially pronounced among women residing in urban areas, and those who are better educated.     

Changes in age at menarche in Spain (1909–1965)

The data used in this work have been collected from clinical histories of women who have been examined at the Ginecology and Obstetritian Service of the Santa cruz y San Pablo Hospital of Barcelona. The sample consisted of 12,803 Spanish women, born between 1909 and 1965, by regions of birth. The method used is retrospective. The maximum menarcheal age in Spain (13.91±0.049 years) is found among those women born between 1925 and 1929 and the minimum menarcheal age (12.92±0.036 years) among women born between 1955 and 1959; so in an interval of 30 years, a reduction of almost a year (11.88 months) has taken place in age at first menstruation. This means a reduction of almost four months per decade. These differences are significant (p=.0000). The higher menarcheal age appears in the Northern Zone, due to the influence of Galicia in the sample (Galicia being a depressed zone with a great emigration rate). The earliest menarcheal age appears in the Mediterranean Zone due to the great percentage of catalan women (Catalonia being a region with a higher socioeconomical level). It is also to be observed that although the menarcheal age has decreased in all the zones, it has done so more dramatically in those zones which had higher values at the beginning. Although the secular trend of reduction in the age at menarche varies greatly from region to region, it may be considered that, in general, the socieconomic conditions have been the decisive factor in the change in physical maturation of Spanish women.     

Current age of the menarche in Belgium

In Belgium, as in most industrialized countries, studies about menarcheal age have shown an earlier physical and sexual maturation. In this paper we will consider the secular trend in menarcheal age over the last twenty years in Brussels. The results concerning menstruation have been obtained in 1980-1982 from 1048 Belgian girls aged between 9 and 17. These girls have very varied socio-economic backgrounds, representative of the diversity of social origin in the Belgian population of Brussels (city and suburbs). Our data were collected using the status quo-method and compared with Jeurissen's (menarcheal age in Belgium between 1915 and 1959). Percentiles and probits were used. The lower limit, where 10% of the girls have had their menarches remains more or less constant. Between 1915 and 1959, Jeurissen noted the tendency of the median menarcheal age toward earlier sexual maturation from 14 years 4 months to 13.0. But, for the first time in Belgium, it seems that the median age is stabilized at 13.0 years. However, the higher limit (age, where 90% have had their menarches) is attained at a lower age: 15.0 for girls born between 1950-1954 and 14.0 in our sample. In conclusion, it seems that better conditions of life cause an evolution in the cases of late menarches but have no more repercussions on the median menarcheal age in this Belgian group.     

Who remains childless?

Who are the men and women who are childless in their mid-30s? Life history data for a British cohort born in 1946 show that age at marriage and marital breakdown were clearly associated with childlessness. Women who were only children were more likely to be childless than those with siblings. Further, early menarcheal age, being highly qualified and having a high status occupation were indirectly related to childlessness. For men, particularly amongst those who had experienced a broken marriage, it was the most ambitious, the highly educated and those in professional occupations who were relatively more likely to be childless.     

LABORATORY EVOLUTION OF LIFE-HISTORY TRAITS IN THE BEAN WEEVIL (ACANTHOSCELIDES OBTECTUS): THE EFFECTS OF DENSITY-DEPENDENT AND AGE-SPECIFIC SELECTION

Four types of laboratory populations of the bean weevil (Acanthoscelides obtectus) have been developed to study the effects of density-dependent and age-specific selection. These populations have been selected at high (K) and low larval densities (r) as well as for reproduction early (Y) and late (O) in life. The results presented here suggest that the r- and K-populations (density-dependent selection regimes) have differentiated from each other with respect to the following life-history traits: egg-to-adult viability at high larval density (K > r), preadult developmental time (r > K), body weight (r > K), late fecundity (K > r), total realized fecundity (r > K), and longevity of males (r > K). It was also found that the following traits responded in statistically significant manner in populations subjected to different age-specific selection regimes: egg-to-adult viability (O > Y), body weight (O > Y), early fecundity (Y > O), late fecundity (O > Y), and longevity of females and males (O > Y). Although several life-history traits (viability, body weight, late fecundity) responded in similar manner to both density-dependent and age-specific selection regimes, it appears that underlying genetic and physiological mechanisms responsible for differentiation of the r/K and Y/O populations are different. We have also tested quantitative genetic basis of the bean weevil life-history traits in the populations experiencing density-dependent and age-specific selection. Among the traits traded-off within age-specific selection regimes, only early fecundity showed directional dominance, whereas late fecundity and longevity data indicated additive inheritance. In contrast to age-specific selecton regimes, three life-history traits (developmental time, body size, total fecundity) in the density-sependent regimes exhibited significant dominance effects. Lastly, we have tested the congruence between short-term and long-term effects of larval densities. The comparisons of the outcomes of the r/K selection regimes and those obtained from the low- and high-larval densities revealed that there is no congruence between the selection results and phenotypic plasticity for the analyzed life-history traits in the bean weevil.     

Genetics of Life History in DROSOPHILA MELANOGASTER. II. Exploratory Selection Experiments

Two types of small-scale selection experiments were performed. (1) Artificial selection experiments were performed on age-specific female fecundity. Selection for early fecundity over three generations produced a statistically detectable direct response. There was no detectable indirect response in other life-history characters. Selection for late fecundity over three generations did not produce any detectable direct response. Indirect responses were detected: early egg-laying decreased and longevity increased. (2) Natural selection for late-age fitness components increased late fecundity, female longevity, and the duration of female reproduction, while early fecundity and mean egg-laying rate decreased.     

Hormonal aspects in the causation of human breast cancer: epidemiological hypotheses reviewed, with special reference to nutritional status and first pregnancy

Epidemiology of breast cancer has identified early age at menarche, late first pregnancy, low parity and late menopause as risk factors, but in addition genetic factors, height, weight and living in western countries play a significant role. The international variation in incidence is almost exclusively due to non-genetic factors. Hypotheses in prevention-oriented research are reviewed: 1. obesity-related oestrogen production as a stimulus of the tumour in postmenopausal women; 2. nutritional status and energy expenditure during puberty and adolescence, developed for fertility and fecundity and extended later to breast cancer; 3. reproductive life during early adulthood, age at first pregnancy and its specific effects on breast tissues. The message of preventability of breast cancer is that mammary epithelial differentiation should come early. Our insight concerning events in puberty and early adulthood can be consolidated in one concept on the risk of extended proliferation of breast epithelium during early adulthood in the absence of full differentiation induced by pregnancy. The combined effects of Western-type nutrition, lack of exercise and Western-type women's emancipation sets the stage for breast cancer already at a young age. Since it is unlikely that emancipated women in affluent societies will return to the original life-style of getting pregnant as soon as it is biologically possible, a novel daring way of protection has to be considered. Could a "Breast Differentiation Pill" be developed to offer protection?     

A Temporal Dimension to the Influence of Pollen Rewards on Bee Behaviour and Fecundity in Aloe tenuior

The net effect of pollen production on fecundity in plants can range from negative – when self-pollen interferes with fecundity due to incompatibility mechanisms, to positive – when pollen availability is associated with increased pollinator visitation and fecundity due to its utilization as a reward. We investigated the responses of bees to pollen and nectar rewards, and the effects of these rewards on pollen deposition and fecundity in the hermaphroditic succulent shrub Aloe tenuior. Self-pollinated plants failed to set fruit, but their ovules were regularly penetrated by self-pollen tubes, which uniformly failed to develop into seeds as expected from ovarian self-incompatibility (or strong early inbreeding depression). Bees consistently foraged for pollen during the morning and early afternoon, but switched to nectar in the late afternoon. As a consequence of this differential foraging, we were able to test the relative contribution to fecundity of pollen- versus nectar-collecting flower visitors. We exposed emasculated and intact flowers in either the morning or late afternoon to foraging bees and showed that emasculation reduced pollen deposition by insects in the morning, but had little effect in the afternoon. Despite the potential for self-pollination to result in ovule discounting due to late-acting self-sterility, fecundity was severely reduced in artificially emasculated plants. Although there were temporal fluctuations in reward preference, most bee visits were for pollen rewards. Therefore the benefit of providing pollen that is accessible to bee foragers outweighs any potential costs to fitness in terms of gender interference in this species.     

Prediction of age at menarche from annual height increments

The divergence in height velocity of early and late maturing girls in early adolescence is exploited to predict menarcheal age from a year's growth in height. Simple prediction equations are generated based on data from the Berkely Guidance Study, and are subsequently tested against an independent sample from the Stuart Growth Study. The success of this prediction method compares favorably with others based on x-ray determinations of bone age; it is much more successful than a proposed method that utilizes estimations of body fat. The height increment method is recommended both by its simplicity and by the relative accessability of the information involved.     

Variable demography in relation to germination time in the annual plant Tagetes micrantha Cav. (Asteraceae)

This study tests whether variability in a life history trait such as emergence time has ecological and evolutionary significance in seasonal environments due to their relationship with fitness components. The population dynamics of nine different cohorts of seedlings emergence were analyzed in a natural population of the annual herb Tagetes micrantha Cav. (Asteraceae). Temporal variation in seedling emergence was related to the pattern of precipitation, while spatial variation could be related to topographical heterogeneity. Plants that emerged at the beginning of the season (early germination) had lower probabilities of survivorship than those that emerged at the end of the season (late germination). In contrast, plants that emerged early had, on average, higher fecundity than those that emerged late. The net result of these contrasting patterns of survival and fecundity constitute a life history trade-off in T. micrantha.     

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

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

Heterosis in age‐specific selected populations of a seed beetle: Sex differences in longevity and reproductive behavior

We tested mutation accumulation hypothesis for the evolution of senescence using short‐lived and long‐lived populations of the seed‐feeding beetle, Acanthoscelides obtectus (Say), obtained by selection on early‐ and late‐life for many generations. The expected consequence of the mutation accumulation hypothesis is that in short‐lived populations, where the force of natural selection is the strongest early in life, the late‐life fitness traits should decline due to genetic drift which increases the frequency of mutations with deleterious effects in later adult stages. Since it is unlikely that identical deleterious mutations will increase in several independent populations, hybrid vigor for late‐life fitness is expected in offspring obtained in crosses among populations selected for early‐life fitness traits. We tested longevity of both sexes, female fecundity and male reproductive behavior for hybrid vigor by comparing hybrid and nonhybrid short‐lived populations. Hybrid vigor was confirmed for male virility, mating speed and copulation duration, and longevity of both sexes at late ages. In contrast to males, the results on female fecundity in short‐lived populations did not support mutation accumulation as a genetic mechanism for the evolution of this trait. Contrary to the prediction of this hypothesis, male mating ability indices and female fecundity in long‐lived populations exhibited hybrid vigor at all assayed age classes. We demonstrate that nonhybrid long‐lived populations diverged randomly regarding female and male reproductive fitness, indicating that sexually antagonistic selection, when accompanied with genetic drift for female fecundity and male virility, might be responsible for overriding natural selection in the independently evolving long‐lived populations.     

Heat shock proteins mediate trade‐offs between early‐life reproduction and late survival in Drosophila melanogaster

Ageing and the resulting increased likelihood mortality are the inescapable fate of organisms because selection pressures on genes that exert their function late in life is weak, promoting the evolution of genes that enhance early‐life reproductive performance at the same time as sacrificing late survival. Heat shock proteins (HSP) are known to buffer various environmental stresses and are also involved in protein homeostasis and longevity. The characteristics of genes for HSPs (hsp) imply that they affect various life‐history traits, which in turn affect longevity; however, little is known about the effects of hsp genes on life‐history traits and their interaction with longevity. In the present study, the effects of hsp genes on multiple fitness traits, such as locomotor activity, total fecundity, early fecundity and survival time, are investigated in Drosophila melanogaster Meigen using RNA interference (RNAi). In egg‐laying females, RNAi knockdown of six hsp genes (hsp22, hsp23, hsp67Ba, hsp67Bb, hsp67Bc and hsp27‐like) does not shorten survival but rather increases it. Knockdown of five of those genes on an individual basis reduces early‐life reproduction, suggesting that several hsp genes mediate the trade‐off between early reproduction and late survival. The data indicate a positive effect of hsp genes on early reproduction and also negative effects on survival time, supporting the antagonistic pleiotropic effects predicted by the optimality theory of ageing.