The variations of human sex ratio at birth with time of conception within the cycle, coital rate around the time of conception, duration of time taken to achieve conception, and duration of gestation: a synthesis

There is a large research literature on the variation of human sex ratio (proportion of males at birth) with (1) time of insemination within the mother's fruitful cycle (TWC), (2) duration of gestation (DOG), (3) coital frequency, here called ‘coital rate’ (CR) and (4) duration of time taken to achieve conception in a period of risk (viz. in the absence of birth limitation methods) (TTC). The variation of sex ratio with each of these four variables has usually been treated as a discrete topic. Consider the four propositions that each of these sorts of variation exists. Here it is argued that these propositions entail one another to varying degrees, and that, for that reason, empirical failures to detect (at conventional levels of significance) one such form of variation (as e.g. with time to conception) should not justify rejecting the hypothesis that such variation exists until the whole network of propositions has been considered. Evidence that offspring sex ratio varies with time of conception within the cycle is strong. It is argued here that, as a consequence, the available data constitute evidence that sex ratio varies with CR and with time to achieve conception, although this variation is small, difficult to detect and of no clinical significance. Lastly, sex ratio varies substantially with DOG, though the explanation for this is not established: it is suggested that the present treatment provides a testable framework for such an explanation.     

Phylogeny of sex‐determining mechanisms in squamate reptiles: are sex chromosomes an evolutionary trap?

Squamate reptiles possess two general modes of sex determination: (1) genotypic sex determination (GSD), where the sex of an individual is determined by sex chromosomes, i.e. by sex‐specific differences in genotype; and (2) temperature‐dependent sex determination (TSD), where sex chromosomes are absent and sex is determined by nongenetic factors. After gathering information about sex‐determining mechanisms for more than 400 species, we employed comparative phylogenetic analyses to reconstruct the evolution of sex determination in Squamata. Our results suggest relative uniformity in sex‐determining mechanisms in the majority of the squamate lineages. Well‐documented variability is found only in dragon lizards (Agamidae) and geckos (Gekkota). Polarity of the sex‐determining mechanisms in outgroups identified TSD as the ancestral mode for Squamata. After extensive review of the literature, we concluded that to date there is no known well‐documented transition from GSD to TSD in reptiles, although transitions in the opposite direction are plentiful and well corroborated by cytogenetic evidence. We postulate that the evolution of sex‐determining mechanisms in Squamata was probably restricted to the transitions from ancestral TSD to GSD. In other words, transitions were from the absence of sex chromosomes to the emergence of sex chromosomes, which have never disappeared and constitute an evolutionary trap. This evolutionary trap hypothesis could change the understanding of phylogenetic conservatism of sex‐determining systems in many large clades such as butterflies, snakes, birds, and mammals. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 168–183.     

A cryptic heterogametic transition revealed by sex-linked DNA markers in Palearctic green toads

In sharp contrast to birds and mammals, most cold‐blooded vertebrates have homomorphic (morphologically undifferentiated) sex chromosomes. This might result either from recurrent X‐Y recombination (occurring e.g. during occasional events of sex reversal) or from frequent turnovers (during which sex‐determining genes are overthrown by new autosomal mutations). Evidence for turnovers is indeed mounting in fish, but very few have so far been documented in amphibians, possibly because of practical difficulties in identifying sex chromosomes. Female heterogamety (ZW) has long been established in Bufo bufo, based on sex reversal and crossing experiments. Here, we investigate a sex‐linked marker identified from a laboratory cross between Palearctic green toads (Bufo viridis subgroup). The F₁ offspring produced by a female Bufo balearicus and a male Bufo siculus were phenotypically sexed, displaying an even sex ratio. A sex‐specific marker detected in highly reproducible AFLP genotypes was cloned. Sequencing revealed a noncoding, microsatellite‐containing fragment. Reamplification and genotyping of families of this and a reciprocal cross showed B. siculus to be male heterogametic (XY) and suggested the same system for B. balearicus. Our results thus reveal a cryptic heterogametic transition within bufonid frogs and help explain patterns of hybrid fitness within the B. viridis subgroup. Turnovers of genetic sex‐determination systems may be more frequent in amphibians than previously thought and thus contribute to the prevalence of homomorphic sex chromosomes in this group.     

Genetic conflict and changes in heterogametic mechanisms of sex determination

The consequences of cytoplasmic sex‐ratio distortion and host repression for the evolution of host sex‐determining mechanisms are examined. Analytical models and simulations are developed to investigate whether the interplay between sex‐ratio distorters and host masculinizers or resistance genes can cause heterogamety switching (changes between male and female heterogamety). Switches from female heterogamety to a system analogous to male heterogamety can occur when selection favours the spread of autosomal masculinizers. However, the evolutionary outcome depends on the type of repressor and costs associated with repression, and also on aspects of population structure. Under most conditions, systems evolved to a polymorphic sex‐determining state although many systems were characterized by numerical dominance of male heterogamety.     

Secondary sex ratios by maternal rank,parity, and age in captive rhesus macaques (Macaca mulatta)

Analysis of 645 conceptions by captive rhesus macaques (Macaca mulatta) with known reproductive histories indicated that maternal age and parity had little effect on the sex of progeny. There was, however, a slight indication that high-ranking females produced a higher proportion of daughters and relatively fewer sons than did low-ranking females. These results provide only weak support for the “advantaged daughter” hypothesis suggested by three previous studies of cercopithecine monkeys, and therefore they further confound the conflicting results for macaques and baboons in general.     

Size distributions and sex ratios of colonizing lizards

This paper reports the body size distributions and sex ratios of four species of phyrnosomatid lizard that colonized experimentally created density sinks. The experiments were conducted in western Colorado in 1992, and lizards colonizing the habitats in 1993 were compared to those removed in 1992 and those present in 1991. Lizards colonizing the density sinks were able to disperse from adjacent habitat. For two of the species (Urosaurus ornatus and Sceloporus undulatus), colonizing lizards were significantly smaller than either those removed in 1992 or those present in 1991. Two other species (S. graciosus and Uta stansburiana) showed no difference in the size distribution of colonizing and removed lizards. In addition, sex ratios of colonizing lizards did not differ from those removed in 1992 or present in 1991. The results of the experiments have implications for the dynamics of the target populations, rescue of local populations from extinction, the regional persistence of populations subject to turnover in patchy environments, and priority effects in colonization. Received: 3 February 1997 / Accepted: 20 May 1998     

Adaptive non-Fisherian sex ratios in a patchily distributed population with outbreeding

A model for sex allocation is presented where mothers have differing abilities to assess their arrival order at a patch in low-density populations. Mothers either oviposit with no knowledge, some knowledge, or they know their arrival order exactly. Furthermore they deposit single eggs in patches hence all matings within a patch are between unrelated individuals. The male offspring however, are also able to mate with females from male-less patches through dispersal. Whereas the first model predicted sex ratios of equality, the models incorporating knowledge both predicted female biased sex ratios. This is shown to be due to an asymmetry in mating opportunities that we have termed random asymmetrical mate competition (RAMC) that only knowledgeable females are able to use to their advantage. Data from the out-breeding non-pollinating fig wasp Otitesella pseudoserrata, suggests that these mothers do have knowledge of their arrival orders and supports the concept and predictions of RAMC.     

Dioecy and the evolution of sex ratios in ants

Split sex ratios, when some colonies produce only male and others only female reproductives, is a common feature of social insects, especially ants. The most widely accepted explanation for split sex ratios was proposed by Boomsma and Grafen, and is driven by conflicts of interest among colonies that vary in relatedness. The predictions of the Boomsma–Grafen model have been confirmed in many cases, but contradicted in several others. We adapt a model for the evolution of dioecy in plants to make predictions about the evolution of split sex ratios in social insects. Reproductive specialization results from the instability of the evolutionarily stable strategy (ESS) sex ratio, and is independent of variation in relatedness. We test predictions of the model with data from a long-term study of harvester ants, and show that it correctly predicts the intermediate sex ratios we observe in our study species. The dioecy model provides a comprehensive framework for sex allocation that is based on the pay-offs to the colony via production of males and females, and is independent of the genetic variation among colonies. However, in populations where the conditions for the Boomsma–Grafen model hold, kin selection will still lead to an association between sex ratio and relatedness.     

The frequency, distribution and reproductive biology of dioecious species in the native flora of Britain and Ireland

The proportion of dioecious species in the flora of the British Isles has been underestimated in the past, partly because the total number of native species was overestimated and partly because some dioecious species were omitted for various reasons. We estimate that, in a total native angiosperm flora of 1377 species and species-equivalents, 4.3% (59 species) are dioecious, a substantially higher percentage than the previous estimates of 2, 3.1 and 3%. If gymnosperms are included, 4.4% (61 of 1380 species of seed plant) are dioecious. A review of published work shows that few of these dioecious species have been well-studied, and that for some little or no information is available about their sex ratios and reproductive biology.     

Extreme sex ratio variation in relation to change in condition around conception

Adaptive theory predicts that mothers would be advantaged by adjusting the sex ratio of their offspring in relation to their offspring's future reproductive success. Studies investigating sex ratio variation in mammals have produced notoriously inconsistent results, although recent studies suggest more consistency if sex ratio variation is related to maternal condition at conception, potentially mediated by changes in circulating glucose level. Consequently, we hypothesized that change in condition might better predict sex ratio variation than condition per se. Here, we investigate sex ratio variation in feral horses (Equus caballus), where sex ratio variation was previously shown to be related to maternal condition at conception. We used condition measures before and after conception to measure the change in condition around conception in individual mothers. The relationship with sex ratio was substantially more extreme than previously reported: 3% of females losing condition gave birth to a son, whereas 80% of those females that were gaining condition gave birth to a son. Change in condition is more predictive of sex ratio than actual condition, supporting previous studies, and shows the most extreme variation in mammals ever reported.     

Sons are made from old stores: sperm storage effects on sex ratio in a lizard

Sperm storage is a widespread phenomenon across taxa and mating systems but its consequences for central fitness parameters, such as sex ratios, has rarely been investigated. In Australian painted dragon lizards (Ctenophorus pictus), we describe elsewhere that male reproductive success via sperm competition is largely an effect of sperm storage. That is, sperm being stored in the female reproductive tract out-compete more recently inseminated sperm in subsequent ovarian cycles. Here we look at the consequences of such sperm storage for sex allocation in the same species, which has genetic sex determination. We show that stored sperm have a 23% higher probability of producing sons than daughters. Thus, shifts in sex ratio, for example over the reproductive season, can partly be explained by different survival of son-producing sperm or some unidentified female mechanism taking effect during prolonged storage.