达乌尔黄鼠冬眠期间体温的变化和冬眠模式

用植入式半导体温度记录元件iButton 记录了达乌尔黄鼠冬眠季节及其前后的体温,分析了其冬眠模式和体温调节特点。结果显示:1)实验室条件下,达乌尔黄鼠冬眠季节长短的个体差异较大,可以分成深冬眠型、
少冬眠型和不冬眠型三种类型;2)达乌尔黄鼠在冬季表现出深冬眠阵(最低体温Tbm in <20℃ ,冬眠阵的持续时间BD >24 h)、短冬眠阵(T_bmin < 20℃ , BD≤24h)和日眠阵(T_bmin ≥20℃ , BD≤24 h)3 种类型,最低体温分别
为2.54℃ ± 0.35℃ 、10.05℃ ± 1.97℃ 和23.09℃ ± 0.40℃ ,彼此之间差异显著。日眠阵阵间产热阶段的最高体温为38.09℃ ±0.17℃ ,高于深冬眠阵(37.31℃ ±0.15℃ )和短冬眠阵(37.22℃ ±0.31℃ ); 3)深冬眠阵和日
眠阵中最低体温均与环境温度显著相关,冬眠过程中的最低体温为-2.43℃ ;4)深冬眠过程中,多数个体可以短时(≤3 h)耐受- 2℃ ～ 0℃ 的低温,激醒或继续维持深冬眠,无致死效应,但长时间(15 h)或过度低温
(- 5℃ 以下)的条件下,深冬眠的达乌尔黄鼠被激醒(70% )或死亡(30% ),不能持续冬眠; 5)入眠前10 d体温日波动幅度显著增加,高于出眠后的日体温波动,且多数个体入眠前出现体温的“试降”。表明,冬眠前
入眠的准备阶段,动物的体温调节已开始发生变化;冬季日眠的调节机制可能与冬眠不同;短时- 2℃ ～ 0℃ 的体温对深冬眠的达乌尔黄鼠无致死效应。     

冬眠和非冬眠状态达乌尔黄鼠肾单位及相关功能因子的比较

冬眠是动物应对冬季低温和食物匮乏的一种生存策略。达乌尔黄鼠 (Spermophilus dauricus) 是典型的贮脂类冬眠动物。为研究冬眠动物肾脏的适应机制,本实验采用组织学、血液生化分析及酶联免疫方法检测了夏季活动期 (7月) 、冬眠期 (12月) 和早春出眠后 (3月) 达乌尔黄鼠肾单位形态学及血清肌酐、尿素和抗利尿激素 (ADH) 的变化,并用qPCR方法检测了肾脏水通道蛋白基因 (AQP1、AQP2和AQP3)、ADH受体 (V2R) 及内皮型一氧化氮合酶基因 (eNOS) 的表达。结果发现,冬眠期和早春出眠期的达乌尔黄鼠肾小球密度、近曲小管和远曲小管的相对管径、皮质部近曲小管数与远曲小管数比值均低于夏季活动期;冬眠期血清肌酐和尿素浓度高于夏季活动期和早春出眠期,ADH浓度及其受体V2R基因表达低于夏季活动期;冬眠期AQP1基因表达高于早春出眠期,AQP3基因表达低于夏季活动期,AQP2基因表达无显著差异;冬眠期eNOS基因表达低于早春出眠期。这些结果表明冬眠的达乌尔黄鼠表现出较低的肾功能;不同时期的水通道蛋白,eNOS及ADH表现出适应性的功能调节。该实验结果丰富了对冬眠动物肾脏适应机制的认识。     

光照黑暗循环条件下达乌尔黄鼠的冬眠模式和能量消耗

为研究冬眠季节的光照条件对贮脂类冬眠动物入眠的影响,在达乌尔黄鼠腹腔埋植体温记录元件iButton,在体重高峰后的下降阶段置于5℃和12L:12D的光照条件下,观察测定其冬眠模式和能量消耗。达乌尔黄鼠冬眠模式出现深冬眠型、少冬眠型和不冬眠型,蛰眠阵包括深冬眠阵、短冬眠阵和日眠阵。不同冬眠阵中最低体温、冬眠阵的持续时间、阵间产热的持续时间、冷却速率和复温速率差异显著;阵间产热的最高体温基本相同。平均每日能量消耗在日眠阵中最高、短冬眠阵中居中、深冬眠阵中最低。入眠时间多集中于黑暗时相,觉醒时间多集中于光照时相。本实验结果提示,在冬眠季节施加光照黑暗循环条件可减少达乌尔黄鼠冬眠的时间,升高阵间最低体温,缩短冬眠阵与阵间产热的持续时间,降低复温速率;增加冬眠期间能量消耗。入眠与觉醒受光照条件影响,具有明显的光暗节律。     

黄鼠(Citellus dauricus)基础代谢率、静止代谢率、化学热调节强度的季节性变化研究

本文测定了达乌尔黄鼠(Citellus dauricus)的耗氧量等指标,从代谢率与代谢调节方面探讨其能量代谢的季节性变化规律。结果表明,黄鼠的中性温度区在春、夏、秋各季有2—3℃的差别。此三个季节的基础代谢率(单位:毫升氧/克·小时,下同)分别为3.6182、2.6396、2.6005。春季显著高于夏季与秋季。在冬眠状态下室温为5℃时静止代谢率最低,为0.4436。在非冬眠各个季节,即0℃至下临界温度范围内静止代谢率与环境温度呈负相关并具线性回归。黄鼠的化学热调节强度较低,特别是在秋季和在冬眠状态下这种能力受到较深程度的抑制。黄鼠在非冬眠期的基础代谢率与体表面积有密切相关,冬眠期两者之间的相关不明显。     

攻毒后的阿拉善黄鼠冬眠生态繁殖与抗体动态观察

阿拉善黄鼠在人工饲养下，每日平均食量为８０－１２０ｇ，春季食量为１００－１５０ｇ，进入冬眠期（呈半睡状态）每日还可食３０－５０ｇ。玫毒后存活黄鼠在室温高的环境中，能越冬和冬眠，但冬眠时间较晚，１２月中旬呈全睡状态的只占３０％，睡眠和醒眠呈交替。体温自３２－３５℃了８－６℃时，呈完全冬眠状态。冬眠期呼吸６－８次／分钟。发情时间比自然条件下提前１－２个月，发情高峰出现在２－４月份（２月份发情率达５５．     

达乌尔黄鼠心肌细胞膜电位的耐寒性

用胞内微电极记录达乌尔黄鼠心室乳头肌细胞膜电位,研究它的耐寒性并附带观察季节及冬眠的影响。76％和55％的心肌标本分别在0℃至—5℃之间可诱发动作电位。3例心肌超冷至-5℃时静息电位(RP)66.6mV,为35℃时的80％,动作电位幅值(APA)60.4mV,为66％,但复温后可完全恢复。RP与APA比最大去极化率(dV/dt_(max))和动作电位时程(ADP)对寒冷有较大的抵抗力。季节对心肌膜电位活动有明显的影响,冬季深眠黄鼠的心肌RP和APA在0℃中显著高于冬季活跃组,提示深眠黄鼠的心肌有较大的耐寒性。这些结果说明冬眠型哺乳动物的心肌细胞膜电位比非冬眠型的有显著较大耐寒性。     

冬眠阵过程中黄鼠脑与血浆阿片肽含量的变化

利用放射免疫测定法,观测了自然冬眠阵过程的常温、入眠、深眠、激醒四个阶段中,达乌尔黄鼠不同脑区及血浆中β－内啡肽（β－ＥＰ）与甲啡肽（ＭＥＫ）的含量。结果表明：黄鼠下丘脑、垂体中的β－内啡肽在入眠过程中增多,在深眠状态下的含量又高于入眠期;海马、血浆中的β－内啡肽含量变动趋势正好相反,在入眠期与深眠期低于常温水平;桥延脑中含量在冬眠阵过程中变化不大。甲啡肽含量在入眠期升高,而在深眠期重新下降,不同     

不同季节和冬眠时相中达乌尔黄鼠视前区温敏神经元特性和下丘脑内NA代谢的变化

比较了不同季节和冬眠时相中达乌尔黄鼠 (Citelleusdauricus)下丘脑内去甲肾上腺素 (noradrenaline ,NA)代谢和视前区 (POA)脑片中各类温敏神经元的比例、温度敏感性、放电活动的临界温度及下限温度 .结果表明 :与夏季动物相比 ,( 1)冬眠各时相中POA温敏神经元的比例和温敏性产生了与冬眠体温调节特性相关的适应性改变 ;( 2 )冬季和冬眠中POA神经元放电的下限温度和温敏神经元活动的临界温度均显著下移 ;( 3 )冬眠中POA神经元对NA反应的敏感性增高 ,冷敏神经元对NA的反应从夏季的抑制型转变为冬眠时的兴奋型 ;( 4)入眠和深冬眠时下丘脑内NA的含量和代谢水平下降 ,出眠时代谢水平升高 .这些变化可能解释动物入眠时主动降低体温和出眠时从深低体温中快速地升温的温度调节机理 .     

达乌尔黄鼠实验室饲养、繁殖及其冬眠阵

为探索实验室条件下达乌尔黄鼠饲养与繁殖的方法及冬眠阵的发生规律,参照野生黄鼠冬眠洞穴的主要生态环境参数,建立人工冬眠屋,采用传统锯末技术记录冬眠阵。结果显示: (1) 处于春季繁殖期的黄鼠应以大鼠饲料为主,辅以少量黄瓜等,夏季活跃期交叉饲喂大鼠饲料与兔饲料,辅以多水的瓜果蔬菜,秋季育肥期以大鼠饲料为主,辅以高脂肪高蛋白的花生、豆类等。(2)雌鼠怀孕期为28 d 左右,哺乳期约一个月,雌鼠每窝产仔4 ～ 8 只,平均5.52 只;初生幼鼠两周内忌换垫料,并避免将异味带入鼠房。(3)黄鼠冬眠期从当年11月下旬至次年3 月上旬,平均93.95 d;冬眠阵睡眠时长平均7. 44 d,阵间激醒时长平均1.36 d,睡眠天数占整个冬眠期的89.9% ;整个冬眠期,黄鼠冬眠阵平均7. 55 个。(4)2009 年秋至2011 年春季,自野外共捕回黄鼠185 只, 存活146 只,存活率78. 9% 。在2006、2009 和2011 年的黄鼠繁殖期,共配对25 对,产仔138 只,成活92 只,成活率为66.7% 。结果表明,野生达乌尔黄鼠可在人工饲养条件下实现繁殖,并可在人工冬眠屋成功冬眠。     

冬眠黄鼠代谢、循环、呼吸系统的变化

本文主要讨论冬眠达乌尔黄鼠的生理生态特征。静止代谢以闭路系统呼吸计测定。心电图以XDH-3型心电图仪记录。 1.冬眠期与活动期黄鼠的耗氧量与环境温度间的关系均呈负相关(表1、图1)。活动期平均每日能量消耗为64.87卡/克,冬眠期则大大降低,仅为0.92卡/克。 2.冬眠期白血球(2427.80个/立方毫米)仅为活动期(6,304.30个/立方毫米)的38.5％;而冬眠期红血球(707.80个/立方毫米)却增加12.7％. 3.冬眠黄鼠呈现周期性呼吸模式,并常出现Cheyrie-Stokes呼吸,每次呼吸暂停时间平均为3—10分钟。 4.冬眠黄鼠心脏活动不规律,心率低,冬眠期平均心率为5.55次/分。     

达乌尔黄鼠入眠准备期的体温、代谢率及能量特征

贮脂类动物在冬眠前大量积累脂肪来准备冬眠,并在入眠时迅速降低体温和代谢率。为探究入眠准备期达乌尔黄鼠体温、代谢率、呼吸商及能量代谢的变化,将其入眠准备期分为育肥期、体重高峰期、育肥后期和冬眠前的试降期,使用植入式半导体温度记录元件iButton、开放式代谢仪和改进的代谢笼,监测其体温、代谢率及呼吸商和能量摄入的变化。结果显示:(1)达乌尔黄鼠体温在冬眠前13 - 34 d 开始下降,远早于冬眠但晚于体重高峰期;体重高峰期体温有降低的趋势,持续时间为1 - 3 d;育肥后期体温显著下降,体温日波动幅度增加。(2)体重高峰期的静止代谢率高于育肥期,育肥后期有降低的趋势,试降期最低。(3)呼吸商在体重高峰期先升高,之后迅速衰减;入眠准备期的能量摄入在体重达高峰期前达到最大值。结果表明,达乌尔黄鼠在入眠准备期,其体温和代谢率已开始降低,能源物质已开始转变;体重高峰期可能是达乌尔黄鼠入眠的一个转折点或启动入眠的开关。     

Circannual control of hibernation by HP complex in the brain

Seasonal hibernation in mammals is under a unique adaptation system that protects organisms from various harmful events, such as lowering of body temperature (Tb), during hibernation. However, the precise factors controlling hibernation remain unknown. We have previously demonstrated a decrease in hibernation-specific protein (HP) complex in the blood of chipmunks during hibernation. Here, HP is identified as a candidate hormone for hibernation. In chipmunks kept in constant cold and darkness, HP is regulated by an individual free-running circannual rhythm that correlates with hibernation. The level of HP complex in the brain increases coincident with the onset of hibernation. Such HP regulation proceeds independently of Tb changes in constant warmth, and Tb decreases only when brain HP is increased in the cold. Blocking brain HP activity using an antibody decreases the duration of hibernation. We suggest that HP, a target of endogenously generated circannual rhythm, carries hormonal signals essential for hibernation to the brain.     

Temperature-independence of circannual variations in circadian rhythms of golden-mantled ground squirrels

In golden-mantled ground squirrels, phase angles of entrainment of circadian locomotor activity to a fixed light-dark cycle differ markedly between subjective summer and winter. A change in ambient temperature affects entrainment only during subjective winter when it also produces pronounced effects on body temperature (Tb). It was previously proposed that variations in Tb are causally related to the circannual rhythm in circadian entrainment. To test this hypothesis, wheel-running activity and Tb were monitored for 12 to 14 months in castrated male ground squirrels housed in a 14:10 LD photocycle at 21 degrees C. Animals were treated with testosterone implants that eliminated hibernation and prevented the marked winter decline in Tb; these squirrels manifested circannual changes in circadian entrainment indistinguishable from those of untreated animals. Both groups exhibited pronounced changes in phase angle and alpha of circadian wheel-running and Tb rhythms. Seasonal variation in Tb is not necessary for circannual changes in circadian organization of golden-mantled ground squirrels.     

The dependence of the rate of heart contractions in the long-tailed Yakutsk suslik Citellus undulatus on environmental temperature]

Decrease of ambient temperature (Ta) leads to the increase of the heart rate (HR) in active ground squirrels C. undulatus by 5.3/min/1 degree C in summer and by 3.8/min/1 degree C in winter. In a hibernation state, the dependence of the HR on Ta was in a good agreement with equation HT = 2.53.exp.(0.1.Ta). On entering into hibernation and on arousal, the HR change outruns the corresponding body temperature (Tb) change by 1.5-2 hours. A maximum HR level (up to 400/min and more) was registered on arousal when Tb reached 17-20 degrees C. A minimal HR level (4-5/min) was observed during hibernation at Ta 2-5 degrees C. The maximum Ta, at witch C. undulatus was hibernating, reached 23-24 degrees C, the HR being 23-25/min.     

Torpor and hibernation in a basal placental mammal, the Lesser Hedgehog Tenrec Echinops telfairi

The patterns of heterothermy were measured in Lesser Hedgehog Tenrecs, Echinops telfairi, under semi-natural conditions in an outdoor enclosure during the austral mid-winter in southwestern Madagascar. The animals were implanted with miniaturized body temperature (Tb) loggers (iButtons) that measured body temperature every 42 min for 2 months (May and June). The tenrecs entered daily torpor on all 60 consecutive days of measurement, that is, on 100% of animal days, with body temperature closely tracking ambient temperature (Ta) during the ambient heating phase. The mean minimum daily Tb of the tenrecs was 18.44 +/- 0.50 degrees C (n = 174, N = 3), and never exceeded 25 degrees C whereas, apart from a few hibernation bouts in one animal, the mean maximum daily Tb was 30.73 +/- 0.15 degrees C (n = 167, N = 3). Thus during winter, tenrecs display the lowest normothermic Tb of all placental mammals. E. telfairi showed afternoon and early evening arousals, but entered torpor before midnight and remained in torpor for 12-18 h each day. One animal hibernated on two occasions for periods of 2-4 days. We consider E. telfairi to be a protoendotherm, and discuss the relevance and potential of these data for testing models on the evolution of endothermy.     

Body temperature patterns and use of torpor in an alpine glirid species,woolly dormouse

Woolly dormice, Dryomys laniger Felten and Storch (Senckenbergiana Biol 49(6):429–435, 1968), are a small (20–30 g), omnivorous (mainly insectivorous), nocturnal glirid species endemic to Turkey. Although woolly dormice have been assumed to hibernate during winter, no information exists on body temperature patterns and use of torpor in the species. In the present study, we aimed to determine body temperature patterns and use of torpor in woolly dormice under controlled laboratory conditions. Accordingly, body temperature (Tb) of woolly dormice was recorded using surgically implanted Thermochron iButtons, small and inexpensive temperature-sensitive data loggers. Woolly dormice exhibited robust, unimodal daily Tb rhythmicity during the euthermic stage before the beginning of hibernation. They displayed short torpor before they began hibernation, although the tendency to enter short torpor was different among individuals. Woolly dormice began hibernation within 1–3 days after exposure to cold and darkness, i.e., on October 22–24, and ended hibernation in the first half of April. Hibernation consisted of a sequence of multiday torpor bouts, interrupted by euthermic intervals. Thus, the patterns of hibernation in woolly dormice were similar to those observed in classical hibernating mammals.     

脑室注射6—羟多巴胺对黄鼠科眠入眠的影响

The effect of forced depletion of brain norepinephrine (NE) on the onset of hibernation was observed in the ground squirrel (Citellus dauricus) by intraventricular injection of 6-hydroxydopamine (6-OHDA). The results showed: (1) Intraventricular injection of 100-200 micrograms 6-OHDA, which depleted 50-60% NE, markedly facilitated the onset of hibernation, i.e. the average induction period for hibernation in the treated animals was significantly shorter than that of the natural hibernating animals. (2) The average total torpor time in the treated animals was longer than that of natural hibernating animals. (3) All hibernating animals treated with 6-OHDA were able to wake up from deep hibernation spontaneously and undergo normal hibernation bouts. The results indicate the decrease of NE system activity in brain is one important factor in triggering the onset of hibernation.     

山蝠和黑斑蛙乳酸脱氨酶同工酶及血糖浓度的季节变化

本工作采用聚丙烯酸胺凝胶电泳等方法,对山蝠（Ｎｙｃｔｕｓｎｏｃｔｕｌａ）和黑斑蛙（Ｒａｎａｎｉｇｒｏｍａｃｕｌａｔａ）在冬眠或蛰伏期与活动期的血清、脑、骨胳肌中的乳酸脱氢酶（ｌａｃｔａｔｅｄｅｈｙｄｒｏｇｅｎａｓｅ,ＬＤＨ）同工酶和血糖浓度进行了比较研究。ＬＤＨ同工酶及血糖浓度表现出明显的季节变化。与活动期的动物相比,在冬眠或蛰伏期中的动物的ＬＤＨ１缺乏、ＬＤＨ５明显增多（Ｐ＜０．００１）,血糖浓度明显升高（Ｐ＜０．００１）。此外,ＬＤＨ同工酶和血糖浓度的季节变化在冬眠动物和蛰伏动物之间还存在着明显的差异。这些结果提示,ＬＤＨ同工酶含量和血糖浓度的季节变化不仅与动物的能量代谢水平相适应,而且显示了冬眠和蛰伏两种生理过程中的可能差异。