Temperature-dependent sex determination

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Temperature-dependent sex determination (TSD) is a system in which the temperature the eggs experience determine the sexes of the organisms that hatch[1]. It is most prevalent and common among amniote vertebrates that are classified under the reptile class[1], but is also used among some birds, such as the Australian Brush-turkey. It differs from the chromosomal sex-determination systems common among vertebrates. It is a type of environmental sex determination (ESD); in other ESD systems, some factors such as population determine the sex of organisms (see Polyphenism).

The eggs are affected by the temperature at which they are incubated during the middle one-third of embryonic development[2]. This critical period of incubation is known as the thermosensitive period (TSP)[3]. The specific time of sex-commitment is known due to several authors resolving histological chronology of sex differentiation in the gonads of turtles with TSD[2].

Types

Within the mechanism, two distinct patterns have been discovered and named Pattern I and Pattern II, with Pattern I further divided into IA and IB. Pattern IA has a single transition zone, where eggs predominantly hatch males if incubated below this temperature zone, and predominantly hatch females if incubated above it. Pattern IB also has a single transition zone, but females are produced below it and males above it. Pattern II has two transition zones, with males dominating at intermediate temperatures and females dominating at both extremes[4]. Very near or at the pivotal temperature of sex determination, mixed sex ratios and, more rarely, intersex individuals are produced[5].

In turtles with TSD, males are generally produced at lower incubation temperatures than females (TSD IA), with this change occurring over a range of temperatures as little as 1-2 °C [5]. At cooler temperatures ranging between 22.5 and 27 degrees Celsius mostly male turtles arise, and at warmer temperatures around 30 degrees Celsius only female turtles arise[6]. In lizards and crocodilians, this pattern is reversed (TSD IB).

Hormones In TSD Systems

Synergism between temperature and hormones has also been identified in these systems. Administering estradiol at male-producing temperatures generates females that are physiologically identical to temperature-produced females[7]. The reverse experiment, males produced at female temperatures, only occurs when a nonaromatizable testosterone or an aromatase inhibitor is administered, indicating that the enzyme responsible for conversion of testosterone to estradiol, aromatase, plays a role in female development[8].

Interestingly, hormones and temperature show signs of acting in the same pathway, in that less hormone is required to produce a sexual shift as the incubation conditions near the pivotal temperature. It has been proposed[9] that temperature acts on genes coding for such steroidogenic enzymes, and testing of homologous GSD pathways has provided a genic starting point[10]. Yet, the genetic sexual determination pathway in TSD turtles is poorly understood and the controlling mechanism for male or female commitment has not been identified.

Advantages Over Genotypic Sex Determination

A 1977 theoretical model (the CharnovBull model)[11][12], predicted that selection should favour TSD over chromosome-based systems when "the developmental environment differentially influences male versus female fitness"[1]; this theoretical model was empirically validated thirty years later[1] but the generality of this hypothesis in reptiles is questioned.
"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" [13]

See also

References

  1. ^ a b c d Warner DA, Shine R (2008). "The adaptive significance of temperature-dependent sex determination in a reptile". Nature. 451 (7178): 566–568. doi:10.1038/nature06519. PMID 18204437. {{cite journal}}: Italic or bold markup not allowed in: |journal= (help); Unknown parameter |DUPLICATE DATA: pmid= ignored (help)
  2. ^ a b Wibbels, T., J.J. Bull, and D. Crews 1991. Chronology and morphology of temperature dependent sex determination. The Journal of Experimental Zoology 260: 371-381
  3. ^ V. Delmas, A.-C. Prevot-Julliard, C. Pieau, M. Girondot 2008 “A mechanistic model of temperature-dependent sex determination in a chelonian: the European pond turtle”. (On-line) Functional Ecology. Accessed April 19, 2008 at http://www.blackwell-synergy.com
  4. ^ Eti, turtles of the world. “Temperature dependent sex determination”. ETI. Accessed April 16, 2008
  5. ^ a b Bull, J. J. 1980. Sex determination in reptiles. Quart. Review of Biology 55: 3-21.
  6. ^ Harding, J. 2002. "Clemmys guttata", Animal Diversity Web. Accessed April 16, 2008
  7. ^ Wibbels, T., J.J. Bull, and D. Crews 1991. Synergism between temperature and estradiol: A common pathway in turtle sex determination. The Journal of Experimental Zoology 260: 130-134.
  8. ^ Crews, D. 1996. Temperature-dependent sex determination: the interplay of steroid hormones and temperature. Zoo. Sci. 13:1-13.
  9. ^ Crews, D. 2003. Sex determination: where environment and genetics meet. Evolution and Development 5: 50-55.
  10. ^ Miller, D., J. Summers, and S. Silber. 2004. Environmental versus genetic sex determination: a possible factor in dinosaur extinction?. Fertility and Sterility 81: 954-964.
  11. ^ Bull JJ, Charnov EL (1977). "Changes in the heterogametic mechanism of sex determination". Heredity. 39 (1): 1–14. doi:10.1038/hdy.1977.38. PMID 268319. {{cite journal}}: Unknown parameter |DUPLICATE DATA: pmid= ignored (help)
  12. ^ Charnov EL, Bull J (1977). "When is sex environmentally determined?". Nature. 266 (5605): 828–830. doi:10.1038/266828a0. PMID 865602. {{cite journal}}: Unknown parameter |DUPLICATE DATA: pmid= ignored (help)
  13. ^ (Nelson, Randy. An Introduction to Behavioral Endocrinology. Sinauer Associates: Massachusetts. 2005. pg 136