中国石龙子母体孕期调温诱导幼体表型: 母体操纵假说的实验检测
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南京师范大学生命科学学院江苏省生物多样性与生物技术重点实验室,南京师范大学生命科学学院江苏省生物多样性与生物技术重点实验室,南京师范大学生命科学学院江苏省生物多样性与生物技术重点实验室,琼州学院生物科学与技术学院海南省两栖爬行动物研究重点实验室,琼州学院

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国家自然科学基金项目(31060064);江苏省高校自然科学基金项目(11KJB180004);海南省重点科技计划项目(ZDXM20110008)


Maternal thermoregulation during gestation affects the phenotype of hatchling Chinese skinks (Eumeces chinensis): testing the maternal manipulation hypothesis
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Nanjing Normal University,,,,Qiongzhou University

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    摘要:

    卵胎生是由卵生繁殖模式通过逐渐增加卵滞留和胚胎在母体子宫内发育的时间进化而来的繁殖模式。有鳞类爬行动物(蜥蜴和蛇)有着较高的繁殖模式多样性,因而是研究卵胎生繁殖模式进化及其适应意义的理想动物模型。至今对于卵胎生进化的选择压力尚无定论,目前有3种关于卵胎生进化的假说受到学者的关注,其中母体操纵假说最受关注但尚未得到充分的检测。研究继1995年母体操纵假说提出之后,以栖息于温带气候环境下的卵生中国石龙子(Eumeces chinensis)为模型动物检测该假说。37条中国石龙子怀卵母体采自浙江丽水市郊。将怀卵母体分置于3个热处理中,其中12条母体提供每日14 h光照时间,13条母体提供10 h光照,其余12条母体没有任何调温机会(体温随室内环境温度而改变)。结果显示:怀卵母体选择体温向下漂移。3种处理下的雌体繁殖特征没有显著差异。长、短调温组下母体产卵时间要早于非调温组母体,但新生卵的胚胎历期没有显著差异。用5种热处理孵化卵,孵化温度分别为:1=室内波动温度孵化;2=27 ℃;3=24-30 ℃;4=22-32 ℃(3和4孵化处理中,孵化箱内的温度每隔1d改变1次,即卵分别在22和24 ℃孵化24 h,随后在32和30 ℃孵化24 h,每2d循环1次直至孵出);5=在实验室后院内模拟石龙子野外巢址孵化。结果显示:孵出幼体的体长、腹长和头部大小(头长和头宽)在3个母体热处理间存在显著差异,其他形态学特征不存在母体热处理间的显著差异;孵化温度以及孵化温度和母体热处理的交互作用对所有的幼体形态学特征均无显著影响。孵化温度以及孵化温度和母体热处理的交互作用对幼体疾跑速和最大持续运动距离无显著影响;但不同的母体热处理显著影响幼体疾跑速和最大持续运动距离。研究结果不仅为"热变异对在一定孵化温度范围内表型无显著变化的物种的幼体表型没有重要的修饰作用"这一假说提供了有力证据,并且支持母体操纵假说的两个主要预测:雌体在孕期通过体温漂变行为提供体内胚胎发育的最适热环境,而由母体调温行为诱导的后代表型的变异将增强后代的适合度。

    Abstract:

    Viviparity (live-bearing) is a reproductive mode that evolves from oviparity through gradual increases in the length of egg retention and thus intrauterine development. Squamate reptiles (lizards, snakes and amphisbaenians) provide a very useful model system for studying the evolutionary transition from oviparity to viviparity and the adaptive significance of this transition, because they exhibit a wide reproductive diversity. Approximately 20% of squamate reptiles are viviparous, and this reproductive mode has evolved far more often in this group of animals than in all other non-mammalian vertebrates combined. The selective forces responsible for evolution of viviparity remain unclear. To date, there are three main hypothesis of evolution of viviparity, cold-climate hypothesis, predictability hypothesis, and maternal manipulation hypothesis. Of these three main hypotheses, the maternal manipulation hypothesis is most widely accepted in scientific community, although empirical studies testing this hypothesis are still limited. Here, we used the Chinese skink Eumeces chinensis, a temperate-climate oviparous scincid lizard, as the model system to test the maternal manipulation hypothesis. Thirty-seven gravid females collected from a population in Lishui, Zhejiang province (eastern China) were maintained in the laboratory under three different thermal conditions until oviposition. Twenty-five females were provided with the basking opportunity, with 12 were allowed to bask for 14-h daily (LT treatment), and 13 for 10-h daily (ST treatment). The remaining 12 females were not provided with the basking opportunity (NTR treatment), and their body temperatures simply tracked changes in ambient environment. Females of different treatments did not differ morphologically. Body temperatures selected by females were measured using a UT325 digital thermometer. We measured each female three times, once on each of three consecutive days, beginng at 15:00 and ended within 2 h on each day. Females selected lower body temperatures and thermoregulated more precisely when gravid, and gravid females selected lower body temperatures than did non-gravid females and males. LT and ST females laid eggs earlier than did NTR females. Females maintained under the three thermal regimes did not differ from each other in any reproductive trait examined, including embryonic stage at oviposition. Six eggs randomly selected from each clutch were assigned to one of the following five treatments: in a room where temperatures varied naturally; in a constant-temperature room at 27 ℃; in two incubators where temperatures varied from 24-30 ℃ and 22-32 ℃, respectively; in the backyard of the laboratory mimicking natural nests. Incubation temperature (IT) affected incubation length, but maternal thermoregulation (MT) and the IT × MT interaction did not. MT affected the phenotype hatchlings, but IT and the IT × MT interaction did not. Hatchlings from eggs produced by LT females were longer in SVL, abdomen length and head length, and ran faster than those produced by ST females; hatchlings from eggs produced by NTR females had the shortest SVLs, abdomen lengths and head lengths, and performed worst in the racetrack. Other hatchling morphological traits examined did not differ among treatments. Our data support the two main predications from the maternal manipulation hypothesis: (1) gravid females shift their body temperature to provide the optimal thermal conditions for embryos; and (2) offspring phenotypes induced by maternal thermoregulation will enhance offspring fitness.

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李宏,周宗师,吴延庆,林隆慧,林炽贤.中国石龙子母体孕期调温诱导幼体表型: 母体操纵假说的实验检测.生态学报,2012,32(23):7255~7263

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