Temperature-dependent Sex Determination - Adaptive Significance

Adaptive Significance

The adaptive significance of TSD is currently not well understood. One possible explanation that TSD is common in amniotes is phylogenetic inertia – TSD is the ancestral condition in this clade and is simply maintained in extant lineages because it is currently adaptively neutral or nearly so. Indeed, recent phylogenetic comparative analyses imply a single origin for TSD in most amniotes around 300 million years, with several more recent independent origins of TSD in squamates and turtles. Consequently, the adaptive significance of TSD in all but the most recent origins of TSD may have been obscured by the passage of deep time, with TSD potentially being maintained in many amniote clades simply because it ‘works’ (i.e. has no overall fitness costs along the lines of the phylogenetic inertia explanation).

Other work centers on a 1977 theoretical model (the Charnov–Bull model), predicted that selection should favour TSD over chromosome-based systems when "the developmental environment differentially influences male versus female fitness"; this theoretical model was empirically validated thirty years later but the generality of this hypothesis in reptiles is questioned. This hypothesis is supported by the persistence of TSD in certain populations of spotted skink (Niveoscincus ocellatus), where it is advantageous to have females early in the season. The warmth early in the season ensures female-biased broods that then have more time to grow and reach maturity and possibly reproduce before they experience their first winter, thereby increasing fitness of the individual.

In support of the Charnov and Bull hypothesis, Warner and Shine (2008) showed confidently that incubation temperature influences males’ reproductive success differently than females in Jacky Dragon lizards (Amphibolurus muricatus) by treating the eggs with chemicals that interfere with steroid hormone biosynthesis. These chemicals block the conversion of testosterone to oestradiol during development so each sex offspring can be produce at all temperatures. They found that hatching temperatures that naturally produce each sex maximized fitness of each sex, which provides the substantial empirical evidence in support of the Charnov & Bull model for reptiles.

An alternative hypothesis of adaptive significance was proposed by Bulmer and Bull in 1982 and supported by the work of Pen (2010). They conjectured that disruptive selection produced by variation in the environment could result in an evolutionary transition from ESD to GSD (Bull, Vogt, and Bulmer, 1982). Pen (2010) addresses evolutionary divergence in SDM’s via natural selection on sex ratios. Studying the spotted skink, a small lizard in Tasmania, he observed that the highland population was not affected by temperature, yet there was a negative correlation between annual temperature and cohort sex ratios in the lowlands. The highlands are colder with a higher magnitude of annual temperature fluctuation and a shorter activity season, delaying maturity, thus GSD is favored so sex ratios are not skewed. However, in the lowlands, temperatures are more constant and a longer activity season allows for favorable conditions for TSD. He concluded that this differentiation in climate causes divergent selection on regulatory elements in the sex-determining network allowing for the emergence of sex chromosomes in the highlands.

"Temperature sex determination could allow the mother to determine the sex of her offspring by varying the temperature of the nest in which her eggs are incubated. However there is no evidence thus far that sex ratio is manipulated by parental care"

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