Transitional range of temperature, pivotal temperatures and thermosensitive stages for sex determination in reptiles

1991 ◽  
Vol 12 (2) ◽  
pp. 169-179 ◽  
Author(s):  
C. Pieau ◽  
N. Mrosovsky

AbstractThe function relating phenotypic sex ratio to incubation temperature in reptiles can vary in a number of ways in addition to simple differences in the temperatures giving 50% of each sex. This paper offers terminology and definitions for describing these relationships. These definitions accomodate interactions between genetic and environmental effects on sexual differentiation, and variability within populations. The paper also discusses the concept of a sensitive stage/period within incubation during which temperature can affect the direction of sexual differentiation. Thermosensitive period has previously been assessed in a variety of ways. A suggestion for a more general way of defining thermosensitive stage/period is made.

The various patterns of environmental sex determination in squamates, chelonians and crocodilians are described. High temperatures produce males in lizards and crocodiles but females in chelonians. Original experiments on the effects of incubation at 30 °C (100% females) or 33 °C (100% males) on development in Alligator mississippiensis are described. These include an investigation of the effect of exposing embryos briefly to a different incubation temperature on the sex ratio at hatching, and a study of the effects of 30 °C and 33 °C on growth and development of alligator embryos and gonads. A 7-day pulse of one temperature on the background of another was insufficient to alter the sex ratio dramatically. Incubation at 33 °C increased the rate of growth and development of alligator embryos. In particular, differentiation of the gonad at 33 °C was enhanced compared with 30 °C. A hypothesis is developed to explain the mechanism of temperature-dependent sex determination (TSD) in crocodilians. The processes of primary sex differentiation are considered to involve exposure to a dose of some male-determining factor during a specific quantum of developmental time during early incubation. The gene that encodes for the male- determining factor is considered to have an optimum temperature (33 °C). Any change in the temperature affects the expression o f this gene and affects the dose or quantum embryos are exposed to. In these cases there is production of females by default. The phylogenetic implications of TSD for crocodilians, and reptiles in particular, are related to the life history of the animal from conception to sexual maturity. Those animals that develop under optimal conditions grow fastest and largest and become male. A general association between the size of an animal and its sex is proposed for several types of vertebrate.


2021 ◽  
Author(s):  
◽  
Nicola J Nelson

<p>Juveniles resulting from artificially induced and incubated eggs are often used to found or augment populations of rare reptiles, but both procedures may compromise the health of hatchlings or their fitness in natural environments. I aimed to test whether these procedures affected size or performance of juvenile tuatara, Sphenodon punctatus, New Zealand reptiles with temperature-dependent sex determination (TSD). Size and performance are phenotypic traits likely to influence fitness and eventual lifetime reproductive success, and are thus important measures of the suitability of artificial induction and incubation techniques for conservation management. I incubated 320 tuatara eggs artificially at 18, 21 and 22ºC; 52% of these were obtained by induction, the remainder were collected from natural nests. An additional 25 natural nests were left intact for investigation of TSD and effects of incubation temperature in nature. Juveniles from all incubation regimes were kept for ten months post-hatching in similar rearing conditions and sexed by laparoscopy. Induced eggs were significantly smaller than naturally laid eggs, and resulted in significantly smaller hatchlings, even when variation among clutches was accounted for. Incubation temperature did not greatly influence size at hatching, but was an important determinant of size by ten months of age; initial egg mass was the most important factor affecting size of hatchlings. Data indicate that TSD occurs in nature. The sex of hatchlings from 21 nests was investigated: 10 nests produced 100% male hatchlings, 4 nests produced 100% female hatchlings, and only 7 nests produced mixed sex ratios which ranged from 11% to 88% males. Sex of juveniles was related to temperature with a larger proportion of males produced in warmer nests. The overall percentage of male hatchlings in natural nests was 64%. Hatching success was 65% from natural nests during the 1998/99 season. Incubation temperatures throughout the year ranged from 2.9 to 34.4ºC. Global warming is likely to skew the hatchling sex ratio towards males if female tuatara are unable to select nest sites according to environmental cues. Evidence from size patterns of tuatara incubated in natural nests supports differential fitness models for the adaptive significance of TSD. The evaluation of artificial incubation as a conservation management tool demonstrated that it is a procedure that benefits conservation as it can be used reliably to produce founders; hatching success was 94% during this study. The sex ratio of artificially incubated juveniles can be easily manipulated; the pivotal temperature lies between 21 and 22ºC. Constant artificial incubation conditions resulted in larger juveniles by ten months of age than those from natural incubation. Naturally incubated juvenile tuatara, however, were faster for their size, their reaction norm to predator stimuli was to run, and they were possibly more aggressive, suggesting naturally incubated juveniles could survive better in nature. No firm conclusions can be reached on the quality of artificially incubated juvenile tuatara because further research will be required to establish the relevance of performance test results in nature and consequences of incubation regimes in the longer term with respect to relative fitness of individuals.</p>


1991 ◽  
Vol 69 (9) ◽  
pp. 2306-2310 ◽  
Author(s):  
Samuel F. Lockwood ◽  
Brenden S. Holland ◽  
John W. Bickham ◽  
Brian G. Hanks ◽  
James J. Bull

Variation in genome size within and among populations of the pond slider, Trachemys scripta, a species with temperature-dependent sex determination, was investigated. Because genome size has been shown to affect developmental rate in various organisms, as does incubation temperature, it was hypothesized that genome size could influence sex determination in species with environmental sex determination. Significant variation in DNA content was found between geographic populations and among clutches. No significant differences in mean genome size were observed among samples incubated at different temperatures or between sexes of turtles hatched at a temperature that yields a mixed sex ratio. Thus, it appears that sex determination in T. scripta is accomplished in the absence of sex-specific and incubation-temperature-specific differences in genome size. Preliminary data from two populations, however, suggest that genome size may be significantly correlated with the threshold incubation temperature at which a mixed sex ratio is produced.


1991 ◽  
Vol 69 (2) ◽  
pp. 530-533 ◽  
Author(s):  
D. O. Conover ◽  
S. B. DeMond

We tested for an effect of temperature during embryonic and larval development on the sex ratio of offspring in two cyprinodontid fishes (Fundulus heteroclitus and Cyprinodon variegatus) having life histories in which temperature-dependent sex determination might be expected to occur. In both species, field collections showed that as young of the year recruited to the population, the sex ratio did not vary over time, nor did it deviate from 1:1. In laboratory experiments, there was no influence of incubation temperature on sex ratio in either species and sex ratios were near unity in all treatments. Although there was no evidence of temperature-dependent sex determination in the populations we studied, this result should be confirmed on other populations before it is generalized to the species level.


1995 ◽  
Vol 73 (11) ◽  
pp. 2091-2097 ◽  
Author(s):  
Vanessa Lewis-Winokur ◽  
Robert M. Winokur

Eggs of captive desert tortoises. Gopherus agassizi, incubated at six temperatures (25, 27, 28, 29, 29.4, and 31 °C) produced 107 specimens. Eggs incubated at 31 °C resulted in a male to female sex ratio of 5:7; all other incubation temperatures resulted in males only. Histological examination of gonads revealed that the testes of newly hatched to 3-month-old individuals showed incomplete and poorly developed seminiferous tubules. Female gonads showed a thickened cortex. Incubation times were longer at lower temperatures. Both hatching success and hatchling survivorship were lower at lower incubation temperatures. We confirm that temperature-dependent sex determination occurs in desert tortoises and that the pivotal temperature is between 31 and 32 °C.


2021 ◽  
Author(s):  
◽  
Nicola J Nelson

<p>Juveniles resulting from artificially induced and incubated eggs are often used to found or augment populations of rare reptiles, but both procedures may compromise the health of hatchlings or their fitness in natural environments. I aimed to test whether these procedures affected size or performance of juvenile tuatara, Sphenodon punctatus, New Zealand reptiles with temperature-dependent sex determination (TSD). Size and performance are phenotypic traits likely to influence fitness and eventual lifetime reproductive success, and are thus important measures of the suitability of artificial induction and incubation techniques for conservation management. I incubated 320 tuatara eggs artificially at 18, 21 and 22ºC; 52% of these were obtained by induction, the remainder were collected from natural nests. An additional 25 natural nests were left intact for investigation of TSD and effects of incubation temperature in nature. Juveniles from all incubation regimes were kept for ten months post-hatching in similar rearing conditions and sexed by laparoscopy. Induced eggs were significantly smaller than naturally laid eggs, and resulted in significantly smaller hatchlings, even when variation among clutches was accounted for. Incubation temperature did not greatly influence size at hatching, but was an important determinant of size by ten months of age; initial egg mass was the most important factor affecting size of hatchlings. Data indicate that TSD occurs in nature. The sex of hatchlings from 21 nests was investigated: 10 nests produced 100% male hatchlings, 4 nests produced 100% female hatchlings, and only 7 nests produced mixed sex ratios which ranged from 11% to 88% males. Sex of juveniles was related to temperature with a larger proportion of males produced in warmer nests. The overall percentage of male hatchlings in natural nests was 64%. Hatching success was 65% from natural nests during the 1998/99 season. Incubation temperatures throughout the year ranged from 2.9 to 34.4ºC. Global warming is likely to skew the hatchling sex ratio towards males if female tuatara are unable to select nest sites according to environmental cues. Evidence from size patterns of tuatara incubated in natural nests supports differential fitness models for the adaptive significance of TSD. The evaluation of artificial incubation as a conservation management tool demonstrated that it is a procedure that benefits conservation as it can be used reliably to produce founders; hatching success was 94% during this study. The sex ratio of artificially incubated juveniles can be easily manipulated; the pivotal temperature lies between 21 and 22ºC. Constant artificial incubation conditions resulted in larger juveniles by ten months of age than those from natural incubation. Naturally incubated juvenile tuatara, however, were faster for their size, their reaction norm to predator stimuli was to run, and they were possibly more aggressive, suggesting naturally incubated juveniles could survive better in nature. No firm conclusions can be reached on the quality of artificially incubated juvenile tuatara because further research will be required to establish the relevance of performance test results in nature and consequences of incubation regimes in the longer term with respect to relative fitness of individuals.</p>


Reproduction ◽  
2015 ◽  
Vol 150 (4) ◽  
pp. 279-287 ◽  
Author(s):  
Jessica A McCoy ◽  
Benjamin B Parrott ◽  
Thomas R Rainwater ◽  
Phillip M Wilkinson ◽  
Louis J Guillette

Despite the widespread occurrence of environmental sex determination (ESD) among vertebrates, our knowledge of the temporal dynamics by which environmental factors act on this process remains limited. In many reptiles, incubation temperature determines sex during a discrete developmental window just prior to and coincident with the differentiation of the gonads. Yet, there is substantial variation in sex ratios among different clutches of eggs incubated at identical temperatures during this period. Here, we test the hypothesis that temperatures experienced prior to the reported thermosensitive period for alligators (Alligator mississippiensis) can impact how the sex determination system responds to thermal cues later in development. Temperature shift experiments on eggs collected from the field within 24 h of oviposition were employed to decouple various maternal influences from thermal effects, and results demonstrate a previously undefined window of thermosensitivity occurring by stage 15 of embryonic development, six stages earlier than previously reported. We also examine the intrasexual expression of several male- and female-biased genes and show that while male-biased genes display no intrasexual differences, ovarian CYP19A1 (aromatase) transcript abundance differs by approximately twofold depending on thermal exposures experienced at early stages of embryonic development. These findings expand our understanding of the ESD in the alligator and provide the rationale for reevaluation of the temporal dynamics of sex determination in other crocodilians.


2019 ◽  
Author(s):  
Lauren Lawson ◽  
Njal Rollinson

AbstractA common reptile conservation strategy involves artificial incubation of embryos and release of hatchlings or juveniles into wild populations. Temperature-dependent sex determination (TSD) occurs in most chelonians, permitting conservation managers to bias sex ratios towards females by incubating embryos at high temperatures, ultimately allowing the introduction of more egg-bearing individuals into populations. Here, we revisit classic sex allocation theory and hypothesize that TSD evolved in some reptile groups (specifically, chelonians and crocodilians) because male fitness is more sensitive to condition (general health, vigor) than female fitness. It follows that males benefit more than females from incubation environments that confer high-quality phenotypes, and hence high-condition individuals. We predict that female-producing temperatures, which comprise relatively high incubation temperatures in chelonians and crocodilians, are relatively stressful for embryos and subsequent life stages. We synthesize data from 28 studies to investigate how constant temperature incubation affects embryonic mortality in chelonians with TSD. We find several lines of evidence suggesting that female-producing temperatures, especially relatively warm temperatures, are more stressful than male-producing temperatures, and we find some evidence that pivotal temperatures (TPiv, the temperature that produces a 1:1 sex ratio) exhibit a correlated evolution with embryonic thermal tolerance. If patterns of temperature-sensitive embryonic mortality are also indicative of chronic thermal stress that occurs post hatching, then conservation programs may benefit from incubating eggs close to species-specific TPivs, thus avoiding high-temperature incubation. Indeed, our models predict that, on average, a sex ratio of more than 75% females can generally be achieved by incubating eggs only 1°C above TPiv. Of equal importance, we provide insight into the enigmatic evolution of TSD in chelonians, by providing support to the hypothesis that TSD evolution is related to the quality of the phenotype conferred by incubation temperature, with males produced in high-quality incubation environments.Lay summaryWe analyze data on embryonic mortality under constant-temperature incubation for 15 species of chelonians with temperature-dependent sex determination (TSD). Mortality is lowest near species-specific pivotal temperatures (Tpiv) but increases rapidly above TPiv, consistent with a theory that explains the adaptive significance of TSD. Conservation managers should incubate embryos near TPiv.


2004 ◽  
Vol 181 (3) ◽  
pp. 367-377 ◽  
Author(s):  
C Pieau ◽  
M Dorizzi

In many species of oviparous reptiles, the first steps of gonadal sex differentiation depend on the incubation temperature of the eggs. Feminization of gonads by exogenous oestrogens at a male-producing temperature and masculinization of gonads by antioestrogens and aromatase inhibitors at a female-producing temperature have irrefutably demonstrated the involvement of oestrogens in ovarian differentiation. Nevertheless, several studies performed on the entire gonad/adrenal/mesonephros complex failed to find differences between male- and female-producing temperatures in oestrogen content, aromatase activity and aromatase gene expression during the thermosensitive period for sex determination. Thus, the key role of aromatase and oestrogens in the first steps of ovarian differentiation has been questioned, and extragonadal organs or tissues, such as adrenal, mesonephros, brain or yolk, were considered as possible targets of temperature and sources of the oestrogens acting on gonadal sex differentiation.In disagreement with this view, experiments and assays carried out on the gonads alone, i.e. separated from the adrenal/mesonephros, provide evidence that the gonads themselves respond to temperature shifts by modifying their sexual differentiation and are the site of aromatase activity and oestrogen synthesis during the thermosensitive period. Oestrogens act locally on both the cortical and the medullary part of the gonad to direct ovarian differentiation. We have concluded that there is no objective reason to search for the implication of other organs in the phenomenon of temperature-dependent sex determination in reptiles. From the comparison with data obtained in other vertebrates, we propose two main directions for future research: to examine how transcription of the aromatase gene is regulated and to identify molecular and cellular targets of oestrogens in gonads during sex differentiation, in species with strict genotypic sex determination and species with temperature-dependent sex determination.


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