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密点麻蜥的能量代谢以及三种沙蜥的繁殖生物学研究
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摘要
本论文从能量代谢和繁殖生物学的角度出发,分两部分研究并探讨了密点麻蜥和三种沙蜥对荒漠环境的适应机制。第一部分以密点麻蜥Eremias multiocellata为研究对象,对其雌性繁殖能量消耗和代谢的种群差异做了详细阐述;第二部分以荒漠沙蜥Phrynocephalus przewalskii、变色沙蜥P. versicolor(?)口叶城沙蜥P. axillaris为研究对象,从形态学和雌性繁殖以及孵化条件对卵孵化率、后代形态和运动能力的影响等方面,研究了生境差异对蜥蜴种间形态和繁殖特征的影响。具体结果如下:
     环境温度的变化对卵胎生蜥蜴密点麻蜥的体温、静止代谢率和雌性繁殖能量消耗有显著影响。密点麻蜥的体温随环境温度的上升而升高,雌性密点麻蜥具有一定的体温调节能力,25℃下密点麻蜥的体温比环境温度稍高,35℃下其体温比环境温度稍低;怀孕密点麻蜥临近生产前用于维持怀孕状态的能量消耗(maintenance costs of pregnancy, MCP)在其能量分配中是相对固定的,在25℃、30℃和35℃下分别为4.219,4.220和4.448mg CO2·min-1,占总能量消耗的19.40%、14.15%和12.32%,在低温时MCP占总能量消耗的比例明显升高。作者认为MCP对于维持卵胎生密点麻蜥的怀孕过程具有重要意义,在不利的环境条件下,怀孕雌性可能通过提高MCP在整体能量分配中的比例来保证怀孕的正常进行。
     温度补偿现象在爬行动物中较为普遍。本文的研究结果表明,在密点麻蜥的两个种群之间,生境较为寒冷的天祝种群其静止代谢率以及心脏、肝脏和骨骼肌的线粒体呼吸控制率(RCR)均显著大于民勤种群,天祝种群心肌中的乳酸脱氢酶(LDH)和己糖激酶(HK)活性以及骨骼肌中的LDH活性也显著高于民勤种群。天祝种群的密点麻蜥经过长期的低温适应,线粒体对氧的利用效率较高,心脏和骨骼肌等组织的糖酵解途径加强,维持了较高的能量代谢水平,这对于在较短的活动时间内加速生长发育、增加个体的生存能力有着重要的意义。
     对荒漠沙蜥、变色沙蜥和叶城沙蜥的繁殖生物学特征研究结果表明三种沙蜥的性二态性和繁殖策略有显著差异,荒漠沙蜥和叶城沙蜥个体大小性二态性显著,表现为成体雄性吻肛长(SVL)大于成体雌性,但变色沙蜥个体大小无显著的两性差异。三种沙蜥的局部形态特征皆与体长呈正相关,雄性的尾长、后肢、头长和头宽均大于雌性;荒漠沙蜥和变色沙蜥成体雄性的腹长小于雌性。作者认为这些性别差异很可能是两性间选择压力的不同造成的。另外,种间的形态差异很可能也与生境差异有关,随荒漠沙蜥、变色沙蜥和叶城沙蜥生境平均气温逐渐降低和降水量的逐渐减少,其体型也有逐渐减小的趋势。
     蜥蜴的形态学特征与其生育力和繁殖策略也紧密相关。荒漠沙蜥、变色沙蜥和叶城沙蜥的部分繁殖特征存在共性,如窝卵数和单枚卵大小呈负相关,窝卵重与雌体SVL呈正相关,窝卵数与母体产后状态呈正相关,说明三个物种在后代的数量和大小之间以及窝卵数与母体产后状态之间均存在权衡。同时,三个物种产卵时间、产后母体体重、窝卵数、窝卵重和相对窝卵重等特征都存在显著的种间差异,随荒漠沙蜥、变色沙蜥和叶城沙蜥生境平均气温的下降和生存环境干旱程度的加剧,其产卵时间逐渐推迟,产后母体体重、卵重和窝卵重也逐渐降低。三个物种采取的生殖策略各不相同,荒漠沙蜥为“小窝大卵”,变色沙蜥为“大窝小卵”,叶城沙蜥为“小窝小卵”;荒漠沙蜥和变色沙蜥可能通过增加窝卵数的大小来增加繁殖输出,而叶城沙蜥则通过增加单枚卵重来增加繁殖输出。
     为了探究温度和湿度对卵孵化和孵出幼体形态特征及运动表现的影响,将三种沙蜥的卵在三个恒定温度(26℃、30℃和34℃)和两个湿度(-440kPa和-620kPa)下进行孵化。结果显示湿度对孵化过程和幼体形态特征几乎没有影响,但孵化温度显著影响三种沙蜥的孵化期、孵化成功率、新生幼体的形态学特征和运动能力。26℃和30℃下荒漠沙蜥和变色沙蜥孵化成功率较高,孵出幼体体型较大、发育较好、且有较好的运动表现,这些个体具有较强的适应性。作者推测30℃可能是荒漠沙蜥和变色沙蜥卵孵化的最适温度,而34℃可能是变色沙蜥卵孵化的临界高温。由于叶城沙蜥卵孵化成功率较低,其卵孵化和幼体特征需要进一步研究。
The maintenance costs of pregnancy (MCP) and energy metabolism of lizard Eremias multiocellata as well as sexual size dimorphism (SSD), reproductive strategies and the effects of incubuting condition on the hatching success, morphological features and locomotor performance of offsprings in three lizards Phrynocephalus przewalskii, P. versicolor and P. axillaris were studied. The mechanisms of their physiological and ecological adaptations to the environment were also discussed in this paper. The concrete results are as follows:
     The body temperature (Tb) and resting metabolic rate (RMR) of female Eremias multiocellata, a viviparous lizard, were measured at25,30and35℃during pregnancy and after parturition to assess energy requirement of reproduction. The results showed that female lizards'Tbs were slightly higher than actual ambient temperature in25and30℃groups, while they were slightly lower than ambient temperature in35℃group. Ambient temperature significantly affected RMR and gestation period of females. Energy requirement was constant in non-pregnant females, while it increased in pregnant females. The maximal estimates of MCP accounted for19.40%,14.15%and12.32%of the total metabolic rate in25,30and35℃group, respectively. The results indicated the MCP was an important component of total energy cost for the lizard E. multiocellata. It is suggested that pregnant females could guarantee the pregnancy at low temperature by increasing the portion of MCP in total energy cost.
     The relationships between temperature and the activities of lactate dehydrogenase (LDH) and hexokinase (HK) and mitochondrial respiration were studied in cardiac muscle, liver and gastrocnemius muscle in two populations of E. multiocellata. The enzymatic activity and mitochondrial respiration were very sensitive to temperature. The Tianzhu population of lizard E. multiocellata which were from colder regions exhibited a relative higher resting metabolic rate and mitochondrial respiration rate than Minqin population in three kinds of tissues. LDH and HK activity of cardiac muscle and LDH activity of gastrocnemius muscle in Tianzhu population were also higher than that of Minqin population. The results indicate that Tianzhu population of E. multiocellata could response to cold acclimation not only by increasing the aerobic enzymatic activity of cardiac and gastrocnemius muscle, but also by increasing maximal oxidative capacities of tissue mitochondria. The metabolic adaptation to cold stress may accelerate development and increase survival capacity of the population.
     The morphological features and female reproductive characteristics were studied in three species of toad-headed lizards, P. przewalskii, P. versicolor and P. axillaris. The results showed that males were the larger sex in P. przewalskii and P. axillaris, whereas female and male adults of P. versicolor did not differ in mean SVL. In P. przewalskii and P. versicolor, females were larger in abdomen length but females of P. axillaris were smaller in abdomen length. Females of three species laid a single clutch of1-6eggs per breeding season. The trade-off between size and number of eggs was evident in all species. The female reproductive traits, clutch size, egg mass and relative clutch mass were differed among species, with females of P. przewalskii laying fewer but larger eggs than did females of P. versicolor in the same SVL. P. axillaris had the smallest body size, and laid relative fewer and smaller eggs, which may related to its cool climate and less precipitation. Our result indicated that climate, habitat and population competition may be the proximate factors affecting the reproductive parameters of three lizards.
     Flexible-shelled eggs of lizard P. przewalskii, P. versicolor and P. versicolor were incubated in different thermal and hydric environments to elicit the effects of incubation environment on hatching success, incubation duration and hatchling phenotypes. The embryogenesis of two species were not sensitive to the hydric environment except P. przewalskii incubated in30℃group. Hatching success was much higher at26and30℃than34℃. Hatchling incubated at26and30℃had longer snout-vent lengths, larger body masses, and better locomotor performance than hatchlings incubated at higher temperature. Compared to P. przewalskii, P. versicolor incubated longer, and had smaller hatchlings, which would benefit more to the species survival living at drier wildlife habitat. We conclude that an incubation temperature of30℃would provide the best balance between developmental rate, hatching success, and posthatch performances, and we suspect the upper egg developmental thresholds for P. versicolor egg incubation may near and slightly higher than34℃. Because of the lower hatching success, egg incubation in P. versicolor should be of more concern in the further.
引文
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