In 2008 researchers Daniel Warner and Richard Shine tested the Charnov-Bull model by conducting experiments on the Jacky dragon (Amphibolurus muricatus), in Australia. Their results showed that temperature-dependent sex determination(TSD) evolved in this species as an adaptation to fluctuating environmental temperatures. The Charnov-Bull model, proposed by Eric Charnov and James Bull in 1977, described the evolution of TSD, although the model was, for many years, untested. Many reptiles and some fish exhibit non-genetic sex determination, in which an embryos' environment can influence the sex of the adult organism. Environmental conditions such as humidity or population density can alter sex in some organisms, and a widespread form of non-genetic sex determination is temperature-dependent sex determination. TSD reveals how embryonic development can contribute to the evolution of physiological processes. Researchers have documented TSD in a wide range of species, and they continue to investigate how such a sex determining system has evolved.
In "Behavioral Thermoregulation by Turtle Embryos," published in Proceedings of the National Academy of Sciences in April, 2011, Wei-Guo Du, Bo Zhao, Ye Chen, and Richard Shine report that turtle embryos can move towards warmer temperatures within the egg when presented with a small, 0.8 degrees Celsius gradient. This behavioral thermoregulation may benefit the embryo's fitness by accelerating the rate of development enough to decrease the incubation period by up to four and a half days. Embryos are generally thought to have little control over their surroundings. This study revealed that embryos may be able to control their developmental environment by modifying their behavior.