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PubMed1998

Melatonin, immunity and cost of reproductive state in male European starlings.

The effects of reproductive condition and exogenous melatonin on immune function were investigated in castrated European starlings, Sturnus vulgaris. Photorefractory and photostimulated starlings exposed to long days were implanted with melatonin or with blank capsules. Photostimulated starlings with blank capsules exhibited reduced splenocyte proliferation in response to the T-cell mitogen, concanavalin A, compared with the other long-day birds. Exogenous melatonin prevented the suppression of immune function by photostimulation. Photorefractory starlings, with or without melatonin implants, exhibited enhanced immune function compared with photostimulated starlings implanted with blanks. This enhancement was not mediated by endogenous melatonin, but appeared to be related to changes in reproductive state. In addition to the traditional costs of reproduction in birds (e.g. raising of young), there may be a cost of the reproductive state of starlings (i.e. whether they are photorefractory or photostimulated). These data are, we believe, the first to indicate a direct effect of reproductive state on immune function that is independent of both photoperiod (i.e., changes in the duration of melatonin secretion) and gonadal steroids.

Animals↗
PubMed2000

Longevity and the costs of reproduction in a historical human population.

It has been argued that the priority that natural selection places on reproduction negatively affects other processes such as longevity and the problem posed by this trade-off underlies the disposable soma theory for the evolution of human ageing. Here we examine the relationship between reproduction and longevity in a historical human population (the Krummhörn, north-west Germany 1720-1870). In our initial analyses, we found no support for the hypothesized negative effects of reproduction on longevity: married women who remained childless lived no longer than women who reproduced and women who had few children lived no longer than women who had many children. However, more detailed analyses in relation to socio-economic class revealed that the extent to which reproduction has an effect on longevity is a function of the level of economic deprivation. We found that, when possible sources of confound were controlled for (e.g. duration of marriage and amount of time spent in fecund marriage), there is an increasingly strong relationship between longevity and reproduction with increasing poverty.

Aged↗
PubMed2000

Inbreeding avoidance and reproductive skew in a cooperative mammal.

In animal social groups, socially subordinate individuals frequently show low reproductive success or completely fail to breed. This suppression of subordinate reproduction is currently typically attributed to control by dominant individuals. However, subordinates in cooperative groups often lack access to unrelated mates, and an alternative possibility is that their reproduction is limited by inbreeding avoidance. Using the eusocial Damaraland mole-rat Cryptomys damarensis, this paper provides the first experimental evidence, to our knowledge, for this explanation. Subordinate, non-breeding female mole-rats were given access to unrelated mates while remaining in the presence of dominant females, and many became reproductively active soon after unrelated males were introduced. Inbreeding avoidance and the availability of unrelated mates provides a plausible and untested explanation for variation in reproductive skew across animal societies.

Aggression↗
PubMed2000

Nuptial gifts and the use of body resources for reproduction in the green-veined white butterfly Pieris napi.

Reproduction in butterflies, as in many holometabolous insects, is usually constrained by the amount of nutrients the animals can collect as juveniles. In polyandric species the females can also supplement their larval-derived reserves with protein-rich donations, so-called nuptial gifts, delivered by the males at mating. Recent findings also indicate that females have access to nitrogen from the histolysis of flight muscles in the thorax. This field study examined how butterflies of the polyandric gift-giving species Pieris napi (Lepidoptera: Pieridae) use body resources in their reproduction and how the male donations affect the females use of stored reserves. The results support earlier studies, indicating that females use resources from the breakdown of thorax muscles to increase their reproductive potential and the results also indicate that males also use thorax material in their reproduction. The study also supports recent findings that the male donation increases the breakdown of body resources and thereby boosts the reproductive output of the female.

Animals↗
PubMed2000

Fitness, reproduction and longevity among European aristocratic and rural Finnish families in the 1700s and 1800s.

The life histories of two socio-economically different groups of humans comprising birth cohorts from the 1700s and 1800s were investigated. It was discovered that fertility selection was greater among European aristocrats and mortality selection greater among rural Finns. The life history of the rural Finns involved shorter female life spans, a considerably longer period of reproduction, higher juvenile mortality, a greater total production of offspring and slightly higher individual fitness. In a comparison of parental cohorts, it was discovered that longevity and progeny survival improved significantly from the 1700s to the 1800s. Out of the three factors investigated, longevity was found to influence reproduction and fitness more than socio-economic group or birth cohort. The reproductive efficacy and fitness of women increased along with their life span. However, reproductive success and fitness were lower among women with the longest life span (over 80 years). Among men, reproductive success improved consistently along with the increase in longevity. When birth intervals were examined, it was discovered that the sex of previous offspring did not influence the interval between births.

Adult↗
PubMed2001

Optimal allocation of reproductive effort: manipulation of offspring number and size in the bank vole.

The number of offspring attaining reproductive age is an important measure of an individual's fitness. However, reproductive success is generally constrained by a trade-off between offspring number and quality. We conducted a factorial experiment in order to study the effects of an artificial enlargement of offspring number and size on the reproductive success of female bank voles (Clethrionomys glareolus). We also studied the effects of the manipulations on growth, survival and reproductive success of the offspring. Potentially confounding effects of varying maternal quality were avoided by cross-fostering. Our results showed that the number of offspring alive in the next breeding season was higher in offspring number manipulation groups, despite their smaller body size at weaning. Offspring size manipulation had no effect on offspring growth or survival. Further, the first litter size of female offspring did not differ between treatments. In conclusion, females may be able to increase the number of offspring reaching reproductive age by producing larger litters, whereas increasing offspring size benefits neither the mother nor the offspring.

Animals↗
PubMed2001

Reproductive suppression in female Damaraland mole-rats Cryptomys damarensis: dominant control or self-restraint?

Colonies of Damaraland mole-rats Cryptomys damarensis exhibit a high reproductive skew. Typically one female breeds and the others are anovulatory. Two models, the dominant control model (DCM) and the self-restraint model (SRM), have been proposed to account for this reproductive suppression. The DCM proposes that suppression is under the control of the dominant breeder and is imposed by mechanisms such as aggression, pheromones and interference with copulation, whereas the SRM does not involve aggression directed towards non-breeders and may function in order to minimize inbreeding. We investigated potential proximate mechanisms involved in the suppression of females in a series of experiments. Socially induced stress through aggression did not appear to be responsible for anovulation. Nor did breeders actively interfere with subordinate copulation. Females were physiologically suppressed when housed in intact colonies. However, as predicted by the DCM, they did not become reproductively active when removed from the presence of breeders. We found no evidence that pheromonal cues block ovulation. We suggest that the SRM is the basic model found in the Damaraland mole-rat and that self-restraint functions in order to minimize inbreeding by restricting reproduction until an unrelated male is present. This would explain the rapid onset of reproductive activation in females when paired with an unrelated male, as demonstrated in this study.

Analysis of Variance↗
PubMed2002

Life-history theory, fertility and reproductive success in humans.

According to life-history theory, any organism that maximizes fitness will face a trade-off between female fertility and offspring survivorship. This trade-off has been demonstrated in a variety of species, but explicit tests in humans have found a positive linear relationship between fitness and fertility. The failure to demonstrate a maximum beyond which additional births cease to enhance fitness is potentially at odds with the view that human fertility behaviour is currently adaptive. Here we report, to our knowledge, the first clear evidence for the predicted nonlinear relationship between female fertility and reproductive success in a human population, the Dogon of Mali, West Africa. The predicted maximum reproductive success of 4.1+/-0.3 surviving offspring was attained at a fertility of 10.5 births. Eighty-three per cent of the women achieved a lifetime fertility level (7-13 births) for which the predicted mean reproductive success was within the confidence limits (3.4 to 4.8) for reproductive success at the optimal fertility level. Child mortality, rather than fertility, was the primary determinant of fitness. Since the Dogon people are farmers, our results do not support the assumptions that: (i) contemporary foragers behave more adaptively than agriculturalists, and (ii) that adaptive fertility behaviour ceased with the Neolithic revolution some 9000 years ago. We also present a new method that avoids common biases in measures of reproductive success.

Adaptation, Physiological↗
PubMed2002

Age-specific survival and reproductive probabilities: evidence for senescence in male fallow deer (Dama dama).

Survival and reproduction are key features in the evolution of life-history strategies. In this study, we use capture-mark-resighting and multi-state models to examine survival senescence and reproductive senescence in six successive cohorts of fallow bucks that were studied for 16 years. We found that the overall age-specific survival probabilities of males were highly variable and the best-fitting model revealed that fallow bucks have four life-history stages: yearling, pre-reproductive, prime-age and senescent. Pre-reproductive males (2 and 3 years old) had the highest survival. Survival declined sharply after the age of 9 years, indicating that senescence had begun. When we considered reproducing and non-reproducing males separately, there was no evidence of senescence in the former, and steadily decreasing survival after the onset of social maturity in the latter. Reproduction probability also declined in older males, and thus we provide very strong evidence of senescence. Reproducers had a greater chance of reproducing again in the following year than non-reproducers. Furthermore, there were differences in the survival probabilities, with reproducers consistently surviving better than non-reproducers. In our study population, reproducers allocate more to the effort to reproduce than non-reproducers. Therefore our results indicate the generally higher phenotypic quality of reproducing males. These results, along with earlier studies on the same population, could indicate positive relationships between fitness correlates.

Age Factors↗
PubMed2003

Female multiple mating behaviour, early reproductive failure and litter size variation in mammals.

Female promiscuity is widespread among mammals, although its function is poorly understood. Recently, much interest has been generated by the hypothesis that female promiscuity, combined with post-copulatory paternity-biasing mechanisms, may function to reduce the costs of reproductive failure resulting from genetic incompatibility. Here, a comparative approach is used to determine if average rates of reproductive failure differ for polytocous mammal species with contrasting levels of female multiple-mating behaviour. After control for phylogeny, promiscuous species were found to have significantly lower rates of early reproductive failure than monogamous and polygynous species, in which females are relatively monandrous. Monandrous females appear to compensate for higher early reproductive failure with increased ova production, and thus produce comparable average litter sizes to those of more promiscuous females. However, there is significantly more variation around the average litter sizes produced by relatively monandrous females. These findings are broadly consistent with predictions of the genetic incompatibility avoidance hypothesis, although it is emphasized that alternative explanations cannot be ruled out on the basis of the comparative evidence presented. Further studies are needed to explore ecological correlates of multiple-mating behaviour, to investigate potential post-copulatory paternity-biasing mechanisms, and to identify the causes of reproductive failure in natural mammal populations.

Animals↗
PubMed2003

Are reproductive skew models evolutionarily stable?

Reproductive skew theory has become a popular way to phrase problems and test hypotheses of social evolution. The diversity of reproductive skew models probably stems from the ease of generating new variations. However, I show that the logical basis of skew models, that is, the way in which group formation is modelled, makes use of hidden assumptions that may be problematical as they are unlikely to be fulfilled in all social systems. I illustrate these problems by re-analysing the basic concessive skew model with staying incentives. First, the model assumes that dispersal is an all-or-nothing response: all subordinates disperse as soon as concessions drop below a certain value. This leads to a discontinuous 'cliff-edge' shape of dominant fitness, and it is not clear that selection will balance a population at such an edge. Second, it is assumed that subordinates have perfect knowledge of their benefits if they stay in the group. I examine the effects of relaxing these two assumptions. Relaxing the first one strengthens reproductive skew theory, but relaxing the latter makes evolutionary stability disappear. In cases where subordinates cannot accurately measure benefits provided by the individual dominant with which they live, so that their behaviour instead evolves as a response to population-wide average benefits, the logic of reproductive skew models does not apply. This warns against too indiscriminate an application of reproductive skew theory to problems in social evolution: for example, transactional models of extra-pair paternity assume perfect knowledge of paternity, which is unlikely to hold true in nature. It is recommended that models specify the mechanisms by which individuals can adjust their behaviour to that of others, and pay attention to changes that occur in evolutionary versus behavioural time.

Animals↗
PubMed2004

Interpreting the von Bertalanffy model of somatic growth in fishes: the cost of reproduction.

We develop a model for somatic growth in fishes that explicitly allows for the energy demand imposed by reproduction. We show that the von Bertalanffy (VB) equation provides a good description of somatic growth after maturity, but not before. We show that the parameters of the VB equation are simple functions of age at maturity and reproductive investment. We use this model to show how the energy demands for both growth and reproduction trade off to determine optimal life-history traits. Assuming that both age at maturity and reproductive investment adapt to variations in adult mortality to maximize lifetime offspring production, our model predicts that: (i) the optimal age of maturity is inversely related to adult mortality rate; (ii) the optimal reproductive effort is approximately equal to adult mortality rate. These predictions are consistent with observed variations in the life-history traits of a large sample of iteroparous freshwater fishes.

Age Factors↗
PubMed2004

Differences in the timing of reproduction between urban and forest European blackbirds (Turdus merula): result of phenotypic flexibility or genetic differences?

Species which have settled in urban environments are exposed to different conditions from their wild conspecifics. A previous comparative study of an urban and a forest-living European blackbird population had revealed a three weeks earlier onset of gonadal growth in urban individuals. These physiological adjustments are either the result of genetic differences that have evolved during the urbanization process, or of phenotypic flexibility resulting from the bird's exposure to the different environmental conditions of town or forest. To identify which of these two mechanisms causes the differences in reproductive timing, hand-reared birds originating from the urban and the forest populations were kept in identical conditions. The substantial differences in the timing of reproduction between urban and forest birds known from the field did not persist under laboratory conditions, indicating that temporal differences in reproductive timing between these two populations are mainly a result of phenotypic flexibility. Nevertheless, urban males initiated plasma luteinizing hormone (LH) secretion and testicular development earlier than forest males in their first reproductive season. Moreover, plasma LH concentration and follicle size declined earlier in urban females than in forest females, suggesting that genetic differences are also involved and might contribute to the variations in the timing of reproduction in the wild.

Animals↗
PubMed2006

Adaptive adjustment of offspring sex ratio and maternal reproductive effort in an iteroparous mammal.

Large mammals in seasonal environments have a pattern of high-reproductive synchrony in spring, but how the timing of reproduction affects resource allocation decisions at different stages of the reproductive cycle remains largely unexplored. By manipulating the timing of conception in reindeer (Rangifer tarandus), we tested how the timing of conception affected sex ratio, gestation length and weight development of mother and offspring. Females that conceived at their first ovulation within the rut had a 60.5% probability of producing a male; in contrast, females that conceived a cycle later had a 31.3% probability of producing a male. Late conceiving females had gestation times that were 10 days shorter and the calves were 0.6 kg (9.2%) lighter at birth and 7.4 kg (14.7%) lighter in autumn. Over the year, female weight changes was similar between the groups suggesting reindeer follow a bet-hedging strategy; reducing the quality of this year's offspring to ensure their own future reproduction and survival. Harvesting is often selective leading to skewed sex ratios and age structure, which may influence the timing of reproduction due to females hesitation to mate with young males. Whenever this hesitation is strong enough to increase the frequency of recycling, harvesting is likely to have profound life history consequences.

Animals↗
PubMed2006

Best squirrels trade a long life for an early reproduction.

Age at primiparity plays a crucial role in population dynamics and life-history evolution. Long-term data on female North American red squirrels were analysed to study the fitness consequences of delaying first reproduction. Early breeders were born earlier, had a higher breeding success and achieved a higher lifetime reproductive success than females who delayed their first reproduction, which suggests a higher quality of early breeders. However, early breeders had similar mass when tagged, and similar number of food caches available at one year of age as late breeders. Nevertheless, we found evidence of survival costs of early primiparity. Early breeders had a lower survival between one and two years of age than late breeders and a lower lifespan. Our study points out that two reproductive tactics co-occurred in this population: a tactic based on early maturity at the cost of a lower survival versus a tactic based on delayed maturity and long lifespan. High quality individuals express the most profitable tactic by breeding early whereas low quality individuals do the best of a bad job by delaying their first reproduction.

Animals↗
PubMed1995

Gibberellins promote vegetative phase change and reproductive maturity in maize.

Postembryonic shoot development in maize (Zea mays L.) is divided into a juvenile vegetative phase, an adult vegetative phase, and a reproductive phase that differ in the expression of many morphological traits. A reduction in the endogenous levels of bioactive gibberellins (GAs) conditioned by any one of the dwarf1, dwarf3, dwarf5, or anther ear1 mutations in maize delays the transition from juvenile vegetative to adult vegetative development and from adult vegetative to reproductive development. Mutant plants cease producing juvenile traits (e.g. epicuticular wax) and begin producing adult traits (e.g. epidermal hairs) later than wild-type plants. They also cease producing leaves and begin producing reproductive structures later than wild-type plants. These mutations greatly enhance most aspects of the phenotype of Teopod1 and Teopod2, suggesting that GAs suppress part but not all of the Teopod phenotype. Application of GA 3 to Teopod2 mutants and Teopod1, dwarf3 double mutants confirms this result. We conclude that GAs act in conjunction with several other factors to promote both vegetative and reproductive maturation but affect different developmental phases unequally. Furthermore, the GAs that regulate vegetative and reproductive maturation, like those responsible for stem elongation, are downstream of GA 20 in the GA biosynthetic pathway.

Gibberellins↗
PubMed2000

Does evolution of iteroparous and semelparous reproduction call for spatially structured systems?

A persistent question in the evolution of life histories is the fitness trade-off between reproducing only once (semelparity) in a lifetime or reproducing repeated times in different seasons (iteroparity). The problem can be formulated into a research agenda by assuming that one reproductive strategy is resident (has already evolved) and by asking whether invasion (evolution) of an alternative reproductive strategy is possible. For a spatially nonstructured system, Bulmer (1994) derived the relationship v + PA < 1 (PA is adult survival; vbS and bS are offspring numbers for iteroparous and semelparous breeding strategies, respectively) at which semelparous population cannot be invaded by an iteroparous mutant. When the inequality is changed to v + PA > 1, invasion of a semelparous mutant is not possible. From the inequalities, it is easy to see that possibilities for evolutionary establishment of a novel reproductive strategy are rather narrow. We extended the evolutionary scenario into a spatially structured system with dispersal linkage among the subunits. In this domain, a rare reproductive strategy can easily invade a population dominated by a resident reproductive strategy. The parameter space enabling invasion is far more generous with spatially structured evolutionary scenarios than in a spatially nonstructured system.

Animals↗
PubMed2000

Global phylogeography of a cryptic copepod species complex and reproductive isolation between genetically proximate "populations".

The copepod Eurytemora affinis has a broad geographic range within the Northern Hemisphere, inhabiting coastal regions of North America, Asia, and Europe. A phylogenetic approach was used to determine levels of genetic differentiation among populations of this species, and interpopulation crosses were performed to determine reproductive compatibility. DNA sequences from two mitochondrial genes, large subunit (16S) rRNA (450 bp) and cytochrome oxidase I (COI, 652 bp), were obtained from 38 populations spanning most of the species range and from two congeneric species, E. americana and E. herdmani. Phylogenetic analysis revealed a polytomy of highly divergent clades with maximum sequence divergences of 10% in 16S rRNA and 19% in COI. A power test (difference of a proportion) revealed that amount of sequence data collected was sufficient for resolving speciation events occurring at intervals greater than 300,000 years, but insufficient for determining whether speciation events were approximately simultaneous. Geographic and genetic distances were not correlated (Mantel's test; r = 0.023, P = 0.25), suggesting that populations had not differentiated through gradual isolation by distance. At finer spatial scales, there was almost no sharing of mtDNA haplotypes among proximate populations, indicating little genetic exchange even between nearby sites. Interpopulation crosses demonstrated reproductive incompatibility among genetically distinct populations, including those that were sympatric. Most notably, two geographically distant (4000 km) but genetically proximate (0.96% 16S, 0.15% COI) populations exhibited asymmetric reproductive isolation at the F2 generation. Large genetic divergences and reproductive isolation indicate that the morphologically conservative E. affinis constitutes a sibling species complex. Reproductive isolation between genetically proximate populations underscores the importance of using multiple measures to examine patterns of speciation.

Animals↗