Proposed role of w chromosome inactivation and the absence of dosage compensation in avian sex determination

Three features of avian sex chromosomes - female heterogamety (ZZ male, ZW female), the apparently inactive state of the W chromosome, and dose-dependent expression of Z-linked genes - are examined in regard to their possible relation to sex determination. It is proposed that the W chromosome is facultatively heterochromatic and that the Z and W chromosomes carry one or more homologous sex-determination genes. The absence of dosage compensation in ZZ embryos, and W inactivation in ZW embryos, would then bring about a 2n(ZZ)-n(ZW) inequality in the effective copy number of such genes. The absence of dosage compensation of Z-linked genes in ZZ embryos is viewed as a means by which two copies of Z-W homologous sex determination genes are kept active to meet the requirements of testis determination. W inactivation may promote ovarian development by reducing the effective copy number of these genes from 2n to n. If there is a W-specific gene for femaleness, spread of heterochromatization to this gene in cells forming the right gonadal primordium may explain the latter's normally undifferentiated state; reversal of heterochromatization may similarly explain the development of the right gonad into a testis following left ovariectomy.

Transposable element-medicated evolution of sex: A population genetic model

For a population made up of individuals capable of sexual as well as asexual modes of reproduction, conditions for the spread of a transposable element are explored using a one-locus, two-haplotype model. The analysis is then extended to include the possibility that the transposable element can modulate the probability of sexual reproduction, thus casting Hickey’s (1982,Genetics 101: 519–531) suggestion in a population genetics framework. The model explicitly includes the cost of sexual reproduction, fitness disadvantage to the transposable element, probability of transposition, and the predisposition for sexual reproduction in the presence and absence of the transposable element. The model predicts several kinds of outcome, including initial frequency dependence and stable polymorphism. More importantly, it is seen that for a wide range of parameter values, the transposable element can go to fixation. Therefore it is able to convert the population from a predominantly asexual to a predominantly sexual mode of reproduction. Viewed in conjunction with recent results implicating short stretches of apparently non-coding DNA in sex determination (McCoubreyet al. 1988,Science 242: 1146–1151), the model hints at the important role this mechanism could have played in the evolution of sexuality.

Male-biased adult sex ratios and their significance for cooperative breeding in dhole, Cuon alpinus, packs

Two free-ranging packs of dholes (Asiatic wild dog, Cuon alpinus) were monitored for a period of 6 yr (Sep. 1990-Sep. 1996) in the Mudumalai sanctuary, southern India. Demographic data on age structure, litter-size, sex ratio and age and sex specific dispersal were collected. Behavioural data on social interactions and reproductive behaviour among pack members were obtained to determine the frequencies of dominant and subordinate behaviours shown by malt: and female pack members and a measure of each male's reproductive access to females. Behaviours displayed by pack members at dens were recorded to determine whether any age- or sex-specific role specialization existed during pup care. Tenures for dominant males and females within the pack were calculated to ascertain the rate of breeding vacancies occurring within packs. Approximate levels of genetic relatedness within packs were determined by studying pedigrees. In most years one study pack had a male-biased adult sex ratio. This was caused by almost twofold higher dispersal of adult females over adult males. A considerable variance existed in the percentage of sub-adults dispersing from the two packs. Differences existed in the frequencies of dominant and subordinate behaviours shown by males. For males, dominance ranks and ranks based on submissive behaviours were not correlated with frequencies of reproductive behaviours. Subordinate males also displayed reproductive behaviours. In packs, dominant males had lower tenures than dominant females indicating that among males breeding vacancies arose more quickly. The litter size was found to be negatively correlated with the age of the breeding female. There were no significant differences across individuals of varying age or sex classes in the display of pup care behaviours. Significant differences did exist among individual adults. Genetic relatedness among packs tended to vary temporally as a consequence of possible mating by subordinate animals and immigration of new males into the pack. In conclusion, it appears that males delay dispersal and cooperate within their natal packs because of the variety of reproductive strategies they could pursue within. A combination of ecological constraints and the difficulties of achieving breeding status within non-natal packs may make early dispersal and independent breeding less beneficial.

A tropical oviparous lizard, Calotes versicolor, exhibiting a potentially novel FMFM pattern of temperature-dependent sex determination

Among squamate reptiles, lizards exhibit an impressive array of sex-determining modes viz. genotypic sex determination, temperature-dependent sex determination, co-occurrence of both these and those that reproduce parthenogenetically. The oviparous lizard, Calotes versicolor, lacks heteromorphic sex chromosomes and there are no reports on homomorphic chromosomes. Earlier studies on this species presented little evidence to the sex-determining mechanism. Here we provide evidences for the potential role played by incubation temperature that has a significant effect (P<0.01) on gonadal sex and sex ratio. The eggs were incubated at 14 different incubation temperatures. Interestingly, 100% males were produced at low (25.5 +/- 0.5 degrees C) as well as high (34 +/- 0.5 degrees C) incubation temperatures and 100% females were produced at low (23.5 +/- 0.5 degrees C) and high (31.5 +/- 0.5 degrees C) temperatures, clearly indicating the occurrence of TSD in this species. Sex ratios of individual clutches did not vary at any of the critical male-producing or female-producing temperatures within as well as across the seasons. However, clutch sex ratios were female- or male-biased at intermediate temperatures. Thermosensitive period occurred during the embryonic stages 3033. Three pivotal temperatures operate producing 1:1 sex ratio. Histology of gonad and accessory reproductive structures provide additional evidence for TSD. The sex-determining pattern, observed for the first time in this species, that neither compares to Pattern I [Ia (MF) and Ib (FM)] nor to Pattern II (FMF), is being referred to as FMFM pattern of TSD. This novel FMFM pattern of sex ratio exhibited by C. versicolor may have an adaptive significance in maintaining sex ratio. J. Exp. Zool. 317:3246, 2012. (c) 2011 Wiley Periodicals, Inc.