Evolution of extraordinary female-biased sex ratios: the optimal schedule of sex ratio in local mate competition

Female-biased sex ratio in local mate competition has been well studied both theoretically and experimentally. However, some experimental data show more female-biased sex ratios than the theoretical predictions by Hamilton [1967. Science 156, 477-488] and its descendants. Here we consider the following two effects: (1) lethal male-male combat and (2) time-dependent control (or schedule) of sex ratio. The former is denoted by a male mortality being an increasing function of the number of males. The optimal schedule is analytically obtained as an evolutionarily stable strategy (ESS) by using Pontrjagin's maximum principle. As a result, an ESS is a schedule where only males are produced first, then the proportion of females are gradually increased, and finally only females are produced. Total sex ratio (sex ratio averaged over the whole reproduction period) is more female-biased than the Hamilton's result if and only if the two effects work together. The bias is stronger when lethal male combat is severer or a reproduction period is longer. When male-male combat is very severe, the sex ratio can be extraordinary female-biased (less than 5%). The model assumptions and the results generally agree with experimental data on Melittobia wasps in which extraordinary female-biased sex ratio is observed. Our study might provide a new basis for the evolution of female-biased sex ratios in local mate competition.     

Intralocus sexual conflict and offspring sex ratio

Males and females frequently have different fitness optima for shared traits, and as a result, genotypes that are high fitness as males are low fitness as females, and vice versa. When this occurs, biasing of offspring sex-ratio to reduce the production of the lower-fitness sex would be advantageous, so that for example, broods produced by high-fitness females should contain fewer sons. We tested for offspring sex-ratio biasing consistent with these predictions in broad-horned flour beetles. We found that in both wild-type beetles and populations subject to artificial selection for high- and low-fitness males, offspring sex ratios were biased in the predicted direction: low-fitness females produced an excess of sons, whereas high-fitness females produced an excess of daughters. Thus, these beetles are able to adaptively bias sex ratio and recoup indirect fitness benefits of mate choice.     

2nd to 4th digit ratio and offspring sex ratio

There is evidence that the ratio of the length of the 2nd and 4th digit (2D:4D) is negatively related to prenatal and adult concentrations of testosterone. It has also been reported that high levels of testosterone at conception in both fathers and mothers are associated with an increased sex ratio (proportion of males at birth). It follows from these observations that low values of 2D:4D may be related to high sex ratio. We present evidence from three populations (English, Spanish and Jamaican) that 2D:4D is negatively related to sex ratio, independent of the sex and ethnicity of the parents.     

A mathematical model on the optimal timing of offspring desertion

We consider the offspring desertion as the optimal strategy for the deserter parent, analyzing a mathematical model for its expected reproductive success. It is shown that the optimality of the offspring desertion significantly depends on the offsprings' birth timing in the mating season, and on the other ecological parameters characterizing the innate nature of considered animals. Especially, the desertion is less likely to occur for the offsprings born in the later period of mating season. It is also implied that the offspring desertion after a partially biparental care would be observable only with a specific condition.     

Production of offspring using current income and reserves: size-number trade-off and optimal offspring size

To analyse the effects of current income on the nature of size-number trade-off and optimal offspring size, we developed a model in which offspring grow by absorbing current income and reserves. The offspring continue to grow while the current income is available or the reserves exist, and they cease to grow when the reserves are depleted and the current income ceases. We showed that the size-number trade-off is nonlinear in the region where the number of offspring is smaller than the critical number and linear in the region where the number of offspring is greater than the critical number. In the former region, the reserves are not depleted by the time the current income ceases and the offspring cease to grow when the reserves are depleted, whereas in the latter region, the reserves are depleted before the current income ceases and the offspring production is completed when the current income ceases. The optimal offspring size is the same as that shown in Sakai and Harada (Evolution 55 (2001) 467) if this optimal size is realized in the region of nonlinear trade-off, whereas the optimal offspring size is the same as that shown in Smith and Fretwell (Am. Natur. 108 (1974) 499) if this optimal size is realized in the region of linear trade-off.     

A within-vector mathematical model of Plasmodium falciparum and implications of incomplete fertilization on optimal gametocyte sex ratio

A mathematical model that simulates the within-vector dynamics of Plasmodium falciparum in an Anopheles mosquito is developed, based on experimental data. The model takes a mosquito's blood meal as input and computes the salivary gland sporozoite load as the final output, a probable measure of mosquito infectivity. Computational model results are consistent with observed results in nature. Sensitivity analysis of the model parameters suggests that reducing the gametocyte density in the blood meal most significantly lowers sporozoite load in the salivary glands and hence mosquito infectivity, and is thus an attractive target for malaria control. The model is used to investigate the implication of incomplete fertilization on optimal gametocyte sex ratio. For a single strain, the transition from complete fertilization to increasingly incomplete fertilization shifts that ratio from 1 to N, where N is the number of viable male gametes produced by a single male gametocyte, towards 1 to 1, which is demonstrated to be the limiting ratio analytically. This ratio is then shown to be an evolutionarily stable strategy as well in the limiting case.     

The influence of a positive correlation between clutch size and offspring fitness on the optimal offspring size

Summary The effect is modeled of a positive relationship between clutch size and offspring fitness on the optimal investment in offspring. In species which meet the assumptions of the model, the model predicts a positive correlation between maternal resource level and offspring size. If larger mothers are able to allocate more resources to offspring, then the model would also predict a positive correlation between maternal size and offspring size when the assumptions of the model are met. Thus, this model may help explain both among and within individual variation in offspring size. When offspring are produced in groups and the number of offspring killed per clutch is limited by predator satiation, offspring in larger clutches may experience a higher probability of survival. Such a life style may be found in animals such as sea turtles. Offspring size is positively correlated with maternal size in some members of this group.     

Ecological and phenological covariates of offspring sex ratio in barn swallows

Non-random sex allocation in relation to parental, ecological and phenological factors has been investigated in several correlational studies of birds, mostly based on few breeding seasons and relatively small sample sizes, which have led to different results. We investigated sex ratio of nestling barn swallows (Hirundo rustica) in relation to adult sex ratio, laying date, clutch size, colony size and meteorological conditions in a sample of 553 broods (>2200 nestlings) during 10 years. At the population level, nestling sex ratio varied among years and deviated from parity in two years. Sex ratio among adults did not predict offspring sex ratio in the current or the following year. At the within-family level, the proportion of sons increased with laying date in large clutches, did not vary among clutches of intermediate size, and tended to decline with laying date in small clutches. Large colonies harbored more sons. The proportion of males increased with temperature during laying whereas the effects of temperature during the pre- or post-laying periods and that of rainfall were non-significant. These patterns of variation of offspring sex ratio did not differ between years. Thus, we identified several potential causal sources of variation in barn swallow offspring sex ratio, including temporal, phenological and ecological factors. The observation of an association of offspring sex with temperature during laying is novel for birds and may be mediated by effects on maternal steroid hormones profile. The ecological and evolutionary implications of present findings are discussed in the light of adaptive sex allocation theory.     

Proteomic analysis of mitochondria from Caenorhabditis elegans

Mitochondria play essential roles in cell physiological processes including energy production, metabolism, ion homeostasis, cell growth, aging and apoptosis. Proteomic strategies have been applied to the study of mitochondria since 1998; these studies have yielded decisive information about the diverse physiological functions of the organelle. As an ideal model biological system, the nematode Caenorhabditis elegans has been widely used in the study of several diseases, such as metabolic diseases and cancer. However, the mitochondrial proteome of C. elegans remains elusive. In this study, we purified mitochondria from C. elegans and performed a comprehensive proteomic analysis using the shotgun proteomic approach. A total of 1117 proteins have been identified with at least two unique peptides. Their physicochemical and functional characteristics, subcellular locations, related biological processes, and associations with human diseases, especially Parkinson's disease, are discussed. An orthology comparison was also performed between C. elegans and four other model organisms for a general depiction of the conservation of mitochondrial proteins during evolution. This study will provide new clues for understanding the role of mitochondria in the physiological and pathological processes of C. elegans.     

Birds bias offspring sex ratio in response to livestock grazing

Livestock grazing, which has a large influence on habitat structure, is associated with the widespread decline of various bird species across the world, yet there are few experimental studies that investigate how grazing pressure influences avian reproduction. We manipulated grazing pressure using a replicated field experiment, and found that the offspring sex ratio of a common upland passerine, the meadow pipit Anthus pratensis, varied significantly between grazing treatments. The proportion of sons was lowest in the ungrazed and intensively grazed treatments and highest in treatments grazed at low intensity (by sheep, or a mixture of sheep and cattle). This response was not related to maternal body condition. These results demonstrate the sensitivity of avian reproductive biology to variation in local conditions, and support growing evidence that too much grazing, or the complete removal of livestock from upland areas, is detrimental for common breeding birds.     

Climate effects on offspring sex ratio in a viviparous lizard

1. Understanding individual and population responses to climate change is emerging as an important challenge. Because many phenotypic traits are sensitive to environmental conditions, directional climate change could significantly alter trait distribution within populations and may generate an evolutionary response. 2. In species with environment-dependent sex determination, climate change may lead to skewed sex ratios at hatching or birth. However, there are virtually no empirical data on the putative link between climatic parameters and sex ratios from natural populations. 3. We monitored a natural population of viviparous lizards with temperature-dependent sex determination (Niveoscincus ocellatus) over seven field seasons. Sex ratios at birth fluctuated significantly among years and closely tracked thermal conditions in the field, with the proportion of male offspring increasing in colder years. 4. This is the first study to demonstrate the effect of local climatic conditions (e.g. temperature) on offspring sex ratio fluctuations in a free-living population of a viviparous ectotherm. A succession of warmer-than-usual years (as predicted under many climate-change scenarios) likely would generate female-biased sex ratios at birth, while an increase in interannual variation (as also predicted under climate change scenarios) could lead to significant fluctuations in cohort sex ratios. If cohort sex ratio bias at birth leads to adult sex ratio bias, long-term directional changes in thermal conditions may have important effects on population dynamics in this species.     

Multiple resources and the optimal balance between size and number of offspring

Summary We extend the classical Smith-Fretwell model for the optimal size of an offspring to the case of allocation of two or more fitness enhancing resources. Unlike the results of the single-resource model, the new model predicts that the optimal allocations will depend on the resource pool sizes. We apply this new model to the problem of carbon and nitrogen allocation to seeds and conclude (1) that the optimal seed size (carbon allocation) should be positively correlated with the ratio of the size of the carbon and nitrogen pools available for investment to offspring (C/N ratio) and (2) that there should be a negative correlation between seed size and absolute seed nitrogen content. These results may account for some of the within- and between-plant variation in resource allocation to seeds.     

Finite populations choose an optimal language

This paper studies the evolution of a proto-language in a finite population under the frequency-dependent Moran process. A proto-language can be seen as a collection of concept-to-sign mappings. An efficient proto-language is a bijective mapping from objects of communication to used signs and vice versa. Based on the comparison of fixation probabilities, a method for deriving conditions of evolutionary stability in a finite population [Nowak et al., 2004. Emergence of cooperation and evolutionary stability in finite populations. Nature 428, 246-650], it is shown that efficient proto-languages are the only strategies that are protected by selection, which means that no mutant strategy can have a fixation probability that is greater than the inverse population size. In passing, the paper provides interesting results about the comparison of fixation probabilities as well as Maynard Smith's notion of evolutionary stability for finite populations [Maynard Smith, 1988. Can a mixed strategy be stable in a finite population? J. Theor. Biol. 130, 247-251] that are generally true for games with a symmetric payoff function.     

Male phenotypic quality influences offspring sex ratio in a polygynous ungulate

Evolutionary models of sex ratio adjustment applied to mammals have ignored that females may gain indirect genetic benefits from their mates. The differential allocation hypothesis (DAH) predicts that females bias the sex ratio of their offspring towards (more costly) males when breeding with an attractive male. We manipulated the number of available males during rut in a polygynous ungulate species, the reindeer (Rangifer tarandus), and found that a doubling of average male mass (and thus male attractiveness) in the breeding herd increased the proportion of male offspring from approximately 40 to 60%. Paternity analysis revealed indeed that males of high phenotypic quality sired more males, consistent with the DAH. This insight has consequences for proper management of large mammal populations. Our study suggests that harvesting, by generating a high proportion of young, small and unattractive mates, affects the secondary sex ratio due to differential allocation effects in females. Sustainable management needs to consider not only the direct demographic changes due to harvest mortality and selection, but also the components related to behavioural ecology and opportunities for female choice.