爬行动物的温度依赖型性别决定

脊椎动物性别决定模式一直是进化生物学领域的热点问题,它对个体发育和自然种群性比组成都具有深刻的影响。性别决定模式根据主要成因可分为基因依赖型性别决定(GSD)和环境依赖型性别决定(ESD)2大类,其中温度依赖型性别决定(TSD)又是ESD中的主要性别决定模式。多数羊膜类脊椎动物具有稳定的GSD模式,而爬行动物的性别决定模式则丰富多样,即使是亲缘关系很近的物种也具有不同的模式。研究者们以爬行动物为模型动物开展了许多关于脊椎动物性别决定方面的工作。本文综述了近年来爬行动物TSD的最新研究进展,回顾了温度和性激素对TSD爬行类动物的影响及其进化适应意义,以及气候变化与TSD爬行类的关系,并提出了今后爬行动物TSD研究的重点。     

爬行动物温度决定性别的现象与机制

性别决定是生物学的一个核心问题。性别决定模式依据决定因素可以分为基因型性别决定(GSD)和环境型性别决定(ESD)2类。温度依赖型性别决定(TSD)是一种特殊的ESD模式,胚胎性别是由发育过程中所经历的温度决定。简要评述了爬行动物TSD与GSD的关系、TSD的类型、TSD的生理和生态调控以及分子机制,归纳了TSD的各种适应性意义假说,并提出今后TSD研究的重点方向。     

龟类温度依赖型性别决定的保护生物学意义

龟类的性别决定方式有基因型性别决定(GSD)和温度依赖型性别决定(TSD)两种类型许多龟类都为TSD型性别决定。研究龟类TSD不但在揭示动物性别决定的进化规律方面具有深远意义,而且在濒危物种保护方面也具有重要的应用价值主要对龟类TSD理论在保护生物学上的重要应用进行了介绍和探讨。     

龟鳖类温度依赖型性别决定机制的研究进展

近几十年来,发现很多爬行动物具有温度依赖型性别决定机制(TSD),即性别分化取决于胚胎发育过程中温度敏感期(TSP)的环境温度高低.龟鳖类存在两种TSD模式,即低温产雄性、高温产雌性的TSD Ⅰ a,低温、高温均产雌性而中间温度产雄性的TSDⅡ.TSD机制在生理水平的作用机制主要受到性腺类固醇激素的控制,温度通过影响芳香化酶和5α-还原酶的活性控制雌、雄激素转化,进而决定了个体的性别分化.在分子水平的研究发现:Sf1、Mis、Sox9、Dax1、Wt1和Dmrt1等基因的表达受到温度的影响,参与了龟鳖类性别分化.介绍了关于TSD的进化意义提出的假说,有待进一步验证.     

温度依赖型性别决定在爬行动物中的研究进展

爬行动物性别决定方式主要有遗传依赖型性别决定(genetic sex determination,GSD)和环境依赖型性别决定(environmental sex determination,ESD),而ESD又以温度依赖型性别决定(temperature sex determination,TSD)为主。研究爬行动物TSD有助于人们弄清楚环境条件对物种表型的影响,从而更好地利用环境条件和遗传基础的共同机制来人为的改善或者诱导具TSD型物种的进化方向,以实现自然和人类的最大利益。该篇综述从母系活动、气候变化(全球气候变暖)、类固醇以及TSD机制四个方面总结了近年来关于爬行动物TSD的最新研究。     

九顶山两栖爬行类动物资源及保护

１９９７．４～１９９８．６在德阳市九顶山区对两栖爬行类运行进行了调查,基本查清了该区两栖爬行类动物的种类,分布及栖息环境,并对加强该区两栖爬行类动物资源保护和合理利用提出了建议。该区有两栖类动物１８种,隶２日６科１０属,爬行类动物１８种,隶１日７科１５属。     

北京松山国家级自然保护区两栖爬行动物资源调查

2009～2010年对北京松山国家级自然保护区两栖爬行类进行了4次调查,调查到两栖爬行类11种,隶属于2目7科。古北界10种,东洋界1种。对比1991年的调查结果,此次调查未发现丽斑麻蜥、赤链蛇、红点锦蛇、团花锦蛇、双斑锦蛇、虎斑颈槽蛇6种爬行类。草地生境两栖爬行类物种多样性最高。中国林蛙(主要分布于溪流)和山地麻蜥(主要分布于岩壁)分别是种群数量最多的两栖类和爬行类物种。在保护实践中可优先保护两栖爬行类物种多样性较高的关键区域,以提高保护成效。     

两栖和爬行类生长及生长激素分泌活动的调节

根据生长内分泌学,综述了近年来两栖和爬行类生长激素（ＧＨ）分泌活动的调节及生长激素在两栖爬行类生长中作用这一研究领域所取得的主要成就和研究进展,研究结果表明：在脊椎动物的进化过程中,ＧＨ的化学结构和功能是相当保守的;ＧＨ对两栖爬行类的生长起促进作用;类胰岛素生长因子（ＩＧＦｓ）也能冰分传递ＧＨ促进两栖类的生长;在两栖爬行类,下丘脑对ＧＨ分泌的调控较哺乳类缺乏特异性,利用外源ＧＨ促进两栖爬行类的生长     

Temperature-dependent sex determination in fish revisited: prevalence, a single sex ratio response pattern, and possible effects of climate change

Background

In gonochoristic vertebrates, sex determination mechanisms can be classified as genotypic (GSD) or temperature-dependent (TSD). Some cases of TSD in fish have been questioned, but the prevalent view is that TSD is very common in this group of animals, with three different response patterns to temperature.

Methodology/Principal Findings

We analyzed field and laboratory data for the 59 fish species where TSD has been explicitly or implicitly claimed so far. For each species, we compiled data on the presence or absence of sex chromosomes and determined if the sex ratio response was obtained within temperatures that the species experiences in the wild. If so, we studied whether this response was statistically significant. We found evidence that many cases of observed sex ratio shifts in response to temperature reveal thermal alterations of an otherwise predominately GSD mechanism rather than the presence of TSD. We also show that in those fish species that actually have TSD, sex ratio response to increasing temperatures invariably results in highly male-biased sex ratios, and that even small changes of just 1–2°C can significantly alter the sex ratio from 1∶1 (males∶females) up to 3∶1 in both freshwater and marine species.

Conclusions/Significance

We demonstrate that TSD in fish is far less widespread than currently believed, suggesting that TSD is clearly the exception in fish sex determination. Further, species with TSD exhibit only one general sex ratio response pattern to temperature. However, the viability of some fish populations with TSD can be compromised through alterations in their sex ratios as a response to temperature fluctuations of the magnitude predicted by climate change.     

Climate change overruns resilience conferred by temperature‐dependent sex determination in sea turtles and threatens their survival

Temperature‐dependent sex determination (TSD) is the predominant form of environmental sex determination (ESD) in reptiles, but the adaptive significance of TSD in this group remains unclear. Additionally, the viability of species with TSD may be compromised as climate gets warmer. We simulated population responses in a turtle with TSD to increasing nest temperatures and compared the results to those of a virtual population with genotypic sex determination (GSD) and fixed sex ratios. Then, we assessed the effectiveness of TSD as a mechanism to maintain populations under climate change scenarios. TSD populations were more resilient to increased nest temperatures and mitigated the negative effects of high temperatures by increasing production of female offspring and therefore, future fecundity. That buffered the negative effect of temperature on the population growth. TSD provides an evolutionary advantage to sea turtles. However, this mechanism was only effective over a range of temperatures and will become inefficient as temperatures rise to levels projected by current climate change models. Projected global warming threatens survival of sea turtles, and the IPCC high gas concentration scenario may result in extirpation of the studied population in 50 years.     

Temperature-Dependent Sex Determination in Sea Turtles in the Context of Climate Change: Uncovering the Adaptive Significance

The adaptive significance of temperature-dependent sex determination (TSD) in reptiles remains unknown decades after TSD was first identified in this group. Concurrently, there is growing concern about the effect that rising temperatures may have on species with TSD, potentially producing extremely biased sex ratios or offspring of only one sex. The current state-of the-art in TSD research on sea turtles is reviewed here and, against current paradigm, it is proposed that TSD provides an advantage under warming climates. By means of coadaptation between early survival and sex ratios, sea turtles are able to maintain populations. When offspring survival declines at high temperatures, the sex that increases future fecundity (females) is produced, increasing resilience to climate warming. TSD could have helped reptiles to survive mass extinctions in the past via this model. Flaws in research on sex determination in sea turtles are also identified and it is suggested that the development of new techniques will revolutionize the field.     

The ecology and evolution of temperature‐dependent reaction norms for sex determination in reptiles: a mechanistic conceptual model

Sex‐determining mechanisms are broadly categorised as being based on either genetic or environmental factors. Vertebrate sex determination exhibits remarkable diversity but displays distinct phylogenetic patterns. While all eutherian mammals possess XY male heterogamety and female heterogamety (ZW) is ubiquitous in birds, poikilothermic vertebrates (fish, amphibians and reptiles) exhibit multiple genetic sex‐determination (GSD) systems as well as environmental sex determination (ESD). Temperature is the factor controlling ESD in reptiles and temperature‐dependent sex determination (TSD) in reptiles has become a focal point in the study of this phenomenon. Current patterns of climate change may cause detrimental skews in the population sex ratios of reptiles exhibiting TSD. Understanding the patterns of variation, both within and among populations and linking such patterns with the selection processes they are associated with, is the central challenge of research aimed at predicting the capacity of populations to adapt to novel conditions. Here we present a conceptual model that innovates by defining an individual reaction norm for sex determination as a range of incubation temperatures. By deconstructing individual reaction norms for TSD and revealing their underlying interacting elements, we offer a conceptual solution that explains how variation among individual reaction norms can be inferred from the pattern of population reaction norms. The model also links environmental variation with the different patterns of TSD and describes the processes from which they may arise. Specific climate scenarios are singled out as eco‐evolutionary traps that may lead to demographic extinction or a transition to either male or female heterogametic GSD. We describe how the conceptual principles can be applied to interpret TSD data and to explain the adaptive capacity of TSD to climate change as well as its limits and the potential applications for conservation and management programs.     

Sex determination and optimal development in the Moorish gecko,Tarentola mauritanica

Under temperature sex determination (TSD), sex is determined by temperature during embryonic development. Depending on ecological and physiological traits and plasticity, TSD species may face demographic collapse due to climate change. In this context, asymmetry in bilateral organisms can be used as a proxy for developmental instability and, therefore, deviations from optimal incubation conditions. Using Tarentola mauritanica gecko as a model, this study aimed first to confirm TSD, its pattern and pivotal temperature, and second to assess the local adaptation of TSD and variation of asymmetry patterns across four populations under different thermal regimes. Eggs were incubated at different temperatures, and hatchlings were sexed and measured. The number of lamellae was counted in adults and hatchlings. Results were compatible with a TSD pattern with males generated at low and females at high incubation temperatures. Estimated pivotal temperature coincided with the temperature producing lower embryonic mortality, evidencing selection towards balanced sex ratios. The temperature of oviposition was conservatively selected by gravid females. Asymmetry patterns found were likely related to nest temperature fluctuations. Overall, the rigidity of TSD may compromise reproductive success, and demographic stability in this species in case thermal nest choice becomes constrained by climate change.     

Phylogeny of sex‐determining mechanisms in squamate reptiles: are sex chromosomes an evolutionary trap?

Squamate reptiles possess two general modes of sex determination: (1) genotypic sex determination (GSD), where the sex of an individual is determined by sex chromosomes, i.e. by sex‐specific differences in genotype; and (2) temperature‐dependent sex determination (TSD), where sex chromosomes are absent and sex is determined by nongenetic factors. After gathering information about sex‐determining mechanisms for more than 400 species, we employed comparative phylogenetic analyses to reconstruct the evolution of sex determination in Squamata. Our results suggest relative uniformity in sex‐determining mechanisms in the majority of the squamate lineages. Well‐documented variability is found only in dragon lizards (Agamidae) and geckos (Gekkota). Polarity of the sex‐determining mechanisms in outgroups identified TSD as the ancestral mode for Squamata. After extensive review of the literature, we concluded that to date there is no known well‐documented transition from GSD to TSD in reptiles, although transitions in the opposite direction are plentiful and well corroborated by cytogenetic evidence. We postulate that the evolution of sex‐determining mechanisms in Squamata was probably restricted to the transitions from ancestral TSD to GSD. In other words, transitions were from the absence of sex chromosomes to the emergence of sex chromosomes, which have never disappeared and constitute an evolutionary trap. This evolutionary trap hypothesis could change the understanding of phylogenetic conservatism of sex‐determining systems in many large clades such as butterflies, snakes, birds, and mammals. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 168–183.     

Pattern and scale of geographic variation in environmental sex determination in the Atlantic silverside,Menidia menidia

The Atlantic silverside, Menidia menidia (Pisces: Atherinidae), exhibits an exceptionally high level of clinal variation in sex determination across its geographic range. Previous work suggested linear changes in the level of temperature‐dependent sex determination (TSD) with increasing latitude. Based on comparisons at 31 sites encompassing the entire species’ range, we find that the change in level of TSD with latitude is instead highly nonlinear. The level of TSD is uniformly high in the south (Florida to New Jersey), then declines rapidly into the northern Gulf of Maine where genotypic sex determination (GSD) predominates and then rebounds to moderate levels of TSD in the northern‐most populations of the Gulf of St. Lawrence. Major latitudinal breakpoints occur in central New Jersey (40^oN) and the northern Gulf of Maine (44^oN). No populations display pure TSD or GSD. Length of the growing season is the likely agent of selection driving variation in TSD with a threshold at 210 days. Because gene flow among populations is high, such distinct patterns of geographic variation in TSD/GSD are likely maintained by contemporary selection thereby demonstrating the adaptive fine tuning of sex determining mechanisms.     

Sympathetic neural responses to 24-hour sleep deprivation in humans: sex differences

Sleep deprivation has been linked to hypertension, and recent evidence suggests that associations between short sleep duration and hypertension are stronger in women. In the present study we hypothesized that 24 h of total sleep deprivation (TSD) would elicit an augmented pressor and sympathetic neural response in women compared with men. Resting heart rate (HR), blood pressure (BP), and muscle sympathetic nerve activity (MSNA) were measured in 30 healthy subjects (age, 22 ± 1; 15 men and 15 women). Relations between spontaneous fluctuations of diastolic arterial pressure and MSNA were used to assess sympathetic baroreflex function. Subjects were studied twice, once after normal sleep and once after TSD (randomized, crossover design). TSD elicited similar increases in systolic, diastolic, and mean BP in men and women (time, P < 0.05; time × sex, P > 0.05). TSD reduced MSNA in men (25 ± 2 to 16 ± 3 bursts/100 heart beats; P = 0.02), but not women. TSD did not alter spontaneous sympathetic or cardiovagal baroreflex sensitivities in either sex. However, TSD shifted the spontaneous sympathetic baroreflex operating point downward and rightward in men only. TSD reduced testosterone in men, and these changes were correlated to changes in resting MSNA (r = 0.59; P = 0.04). Resting HR, respiratory rate, and estradiol were not altered by TSD in either sex. In conclusion, TSD-induced hypertension occurs in both sexes, but only men demonstrate altered resting MSNA. The sex differences in MSNA are associated with sex differences in sympathetic baroreflex function (i.e., operating point) and testosterone. These findings may help explain why associations between sleep deprivation and hypertension appear to be sex dependent.