The role of prostaglandins and the hypothalamus in thermoregulation in the lizard, Phrynocephalus przewalskii (Agamidae)

Typically, small lizards rely heavily on behavioral thermoregulation rather than physiological mechanisms to control their rates of warming and cooling. We tested the hypothesis that prostaglandins participate in mediating the cardiovascular response to heating and cooling and temperature regulating neurons in the hypothalamus of the small lizard Phrynocephalus przewalskii. In vivo and in vitro treatments, heart rates (HRs) were all found to be higher during heating than during cooling, hysteresis was distinct below 30 and 26°C, respectively. In vivo, as administration of COX inhibitor, there were no differences in HR between heating and cooling at any body temperature and administration of agonist prostaglandins only produced a significant effect on HR below 25°C. Single-unit activity was recorded extracellularly in vitro with microelectrodes, found the firing rate of the continuous unit increased 23% when the temperature of the artificial cerebrospinal fluid dropped from 30–20°C. We conclude that prostaglandins appear to play only a limited role in modulating heart activity in Phrynocephalus przewalskii and suggest that cold-sensitive neurons in the preoptic and anterior hypothalamus (PO/AH) are involved in thermoregulatory control during heating or cooling.     

Habitat use and activity influence thermoregulation in a tropical lizard, Ameiva exsul

We examined the contributions of alterations in daily activity and behavioral selection of microhabitat to thermoregulation in a population of the lizard, Ameiva exsul (Teiidae), by combining data on lizard activity with data on the availability of sun-shade patches and operative temperatures (T_e). By comparing T_e distributions predicted by “no thermoregulation” and “only thermoregulation” hypotheses to those predicted by random use of thermal habitat, we assessed the relative contributions of microhabitat selection and daily activity to regulation of body temperature (T_b). Over the course of a day lizards maintained T_b very close to optimal temperature (T_sel) despite T_es that deviated substantially from T_sel. Data demonstrating a unimodal daily activity pattern reject the hypothesis of uniform activity throughout the day. Also, lizard activity was not positively correlated with the proportion of T_es within T_sel nor negatively correlated with the absolute deviation of available T_e from T_sel (d_e) (“only thermoregulation”). Microsite use by A. exsul deviated significantly from predictions of the “no thermoregulation” hypothesis, but our data could not reject predictions of the “only thermoregulation” hypothesis that lizards would use sun-shade patches relative to the proportion of microsites where T_e is within T_sel. Also, lizards appeared to actively select sunlit and partially shaded microsites at different times of day. Thus, despite thermal constraints imposed by the habitat, A. exsul maintained high and relatively constant T_bs throughout its daily activity period and thermoregulated effectively. This appears to be generally representative of West Indian species of Ameiva.     

广东罗坑自然保护区鳄蜥的体温调节及静止代谢率的热依赖性

探讨鳄蜥（Shinisaurus crocodilurus）的体温调节和静止代谢率及其热依赖性特征,测定了在自然和实验室条件下的环境温度、鳄蜥体温以及静止代谢率（RMR）等热生物学指标。结果显示：在自然条件下,鳄蜥的野外活动体温（Th）稍高于基质温度（Tc）和空气温度（Td）并与后二者呈正相关关系（Tb=11．65＋0．47Tc,r^2=0．34,F1.75=39．11,P〈0．0001;Tb=11．61＋0．46Td,r^2=0．56,F1.75=96．56,P〈0．0001）。鳄蜥在野外的活动体温存在明显的月份间差异,但无性别及年龄上的差异。环境温度为15℃-30℃的实验条件下,鳄蜥的静止体温（Tb）与环境温度（Ta）呈正相关关系：Tb=12．450＋0．5641Ta（F1.110=11．34,r^2=0．997,P〈0．001）;在缺乏温度梯度的环境中,尽管鳄蜥的体温、空气温度和基质温度都有显著的昼夜差异,但体温、空气温度和基质温度的日平均温度之间无显著差异。鳄蜥的RMR不存在雌雄个体间的差异,并与环境温度呈正相关关系。分析显示鳄蜥的体温既有行为调节也有生理调节,其体温调节及RMR符合变温动物的热生物学特征,但是相对于其他活动较多的蜥蜴而言,鳄蜥的生理调节能力较强,这可能与其活动时间分配中静栖占绝大部分的行为特点是相适应的[动物学报54（6）：964-971,2008]。     

Thermal responses to feeding in a secretive and specialized predator (Gila monster, Heloderma suspectum)

We investigate how a unique dietary specialist, the Gila monster (Heloderma suspectum), uses behavioral thermoregulation to elevate body temperature (T_b) after feeding. Lizards in a laboratory thermal gradient were fed rodent meals of three different sizes (5, 10, or 20% of body mass), or sham fed (meal of 0% body mass), and T_bs were recorded for three days before feeding and seven days after feeding. Gila monsters selected a mean T_b of 25.2 °C while fasting (set-point range 23.6–27.1), and increased T_bs after feeding. The magnitude and duration of post-prandial T_b increases are positively related to meal size, and Gila monsters selected mean T_bs up to 3.0 °C higher and maintain elevated T_bs for 3–6 days after feeding. Selection of T_b does not appear to differ between day and night time periods, and because the lizards are both diurnal and nocturnal (at different times of year), photoperiod may not be an important influence on T_b selection.     

Daily and annual patterns of thermoregulation in painted turtles (Chrysemys picta marginata) living in a thermally variable marsh in Northern Michigan

The capacity for an ectothermic reptile to thermoregulate has implications for many components of its life history. Over two years, we studied thermoregulation in a population of Midland painted turtles (Chrysemys picta marginata) in a shallow, thermally variable wetland during summer in Northern Michigan. Mean body temperature (T_b) of free-ranging turtles was greater in 2008 (25.8 °C) than in 2010 (19.7 °C). Laboratory determined thermoregulatory set point (T_set) ranged from 25 °C (T_set-min) to 31 °C (T_set-max) and was lower during the fall (17–26 °C). Deviations of T_b distributions from field measured operative temperatures (T_e) and indices of thermoregulation indicated that C. picta marginata were capable of a limited degree of thermoregulation. Operative temperatures and thermal quality (d_e=|T_set-min−T_e| and |T_e−T_set-max|) cycled daily with maximal thermal quality occurring during late morning and late afternoon. The accuracy of thermoregulation (d_b=|T_set-min−T_b| and |T_b−T_set-max|) was maximal (d_b values were minimal) as T_b declined and traversed T_set during the late afternoon–early evening hours and was higher on cloudy days than on sunny days because relatively low T_e values decreased the number of T_b values that were above T_set. Our index of thermal exploitation (E_x=frequency of T_b observations within T_set) was 36%, slightly lower than that reported for an Ontario population of C. picta marginata. Regression of d_b (thermal accuracy) on d_e (thermal quality) indicated that turtles invested more in thermoregulation when thermal quality was low and when water levels were high than when they were low. There were no intersexual differences in mean T_b throughout the year but females had relatively high laboratory determined T_b values in the fall, perhaps reflecting the importance of maintaining ovarian development prior to winter.     

Thyroxine affects behavioral thermoregulation but not growth rate among populations of the western fence lizard (Sceloporus occidentalis)

We investigated whether thyroxine influences hatchling growth rate of the western fence lizard (Sceloporus occidentalis) throught its effects on thermoregulatory behaviors. We reared control and thyroxine-injected hatchlings from three populations of S. occidentalis that differ in growth rate in a thermal gradient. We also measured the daily changes in body temperature and activity level (proportion of time spent out of retreat sites) of control and thyroxine-injected lizards. Previous studies have shown that within and among population differences in growth rate of the western fence lizard are correlated with the maintenance of high activity levels (proportion of time spent outside of retreat sites) and high body temperatures throughout the day. Growth rate was not influenced by injections of thyroxine. However, injections of thyroxine did elevate average daily body temperature and daily activity. Although administration of thyroxine uniformly increased the probability of activity throughout the day, it did not appear to alter the daily changes in activity. Previous studies have shown that the slower-growing hatchlings from northern populations exhibit a decline in activity during the later part of the thermal cycle, whereas the faster growing southern hatchlings maintain the same level of high activity throughout the thermal cycle. The decline in activity of northern populations was not prevented by thyroxine injection used in our current study. Northern lizards receiving exogenous thyroxine were still less active later in the day compared to early in the day, even though activity level throughout the day was elevated. Thus, the effects of thyroxine on temperature regulation observed in our study (general increase in activity level) appear to be unrelated to those aspects of temperature regulation (e.g., daily changes in behavioral thermoregulation) that are correlated with among population differences in growth rate. We also raised hatchlings in a cycling thermal regime (forced thermal cycle of 34°C:15°C, 12L:12D) where behavioral thermoregulation is not possible. The growth rate of lizards forced to cycle between 34°C:15°C on a daily basis was significantly lower than those lizards allowed to behaviorally thermoregulate, further underscoring the importance of the circadian pattern of thermoregulation for growth.Abbreviations GR growth rate - MR metabolic rate - SMR standard metabolic rate - SVL snout-vent length - T₄ thyroxine - T _b body temperature - T _e environmental temperature     

Body temperature and thermoregulation of Komodo dragons in the field

Komodo dragons from hatchlings (≈0.1 kg) to adults (≤80 kg) express the full magnitude of varanid species size distributions. We found that all size groups of dragons regulated a similar preferred body temperature by exploiting a heterogeneous thermal environment within savanna, forest and mangrove habitats. All dragons studied, regardless of size, were able to regulate a daytime active body temperature within the range 34–35.6 °C for 5.1–5.6 h/day. The index of effectiveness of thermoregulation (a numerical rating of thermoregulatory activity) was not different among size groups of dragons. However, the index of closeness of thermoregulation, which rates the variability of body temperature, suggests a greater precision for regulating a preferred body temperature for medium compared to small and large dragons. Reference copper cylinders simulating small, medium and large Komodo dragons heated and cooled at the same rate, whereas actual dragons of all size groups heated faster than they cooled. Larger dragons heated and cooled more slowly than smaller ones. The mean operative environmental temperatures of copper cylinders representing medium sized dragons were 42.5, 32.0 and 29.4° C for savannah, forest and mangrove habitats, respectively. The index for average thermal quality of a habitat as measured by the absolute difference between operative environmental temperature and the dragon’s thermal range suggests the forest habitat offers the highest thermal quality to dragons and the savannah the lowest. The percent of total daytime that the operative environmental temperature was within the central 50% of the body temperatures selected by dragons in a thermal gradient (Phillips, 1984) was 45%, 15%, and 9% for forest, mangrove and savannah, respectively. Forest habitat offers the most suitable thermal environment and provides the greatest number of hours with conditions falling within the dragon’s thermal activity zone.     

Seasonal and geographic variation in thermal biology of the lizard Microlophus atacamensis (Squamata: Tropiduridae)

1.

We studied physiological thermal adaptation in the lizard Microlophus atacamensis along a latitudinal range.     

Thermoregulatory movement patterns of the lizard Podarcis carbonelli (Lacertilia: Lacertidae)

Using video cameras and motion detection software, we examined sequential positions of the lizard Podarcis carbonelli in a temperature gradient to look for patterns in spatial and temporal thermoregulatory movements. As lizards shuttled between warm and cool areas, their movements were typically slow; punctuated by bursts of speed. The animals were relatively inactive when heating, moved almost continually when cooling, and spent less time heating than cooling. Traditional modeling techniques proved unsuccessful, so we assessed the movement patterns with nonlinear dynamical techniques. The shuttling frequency, and the pattern of velocity changes, both met the qualitative attributes (self similarity, strange attractors, and noisy power spectra) and the quantitative criteria (positive Lyapunov exponent and capacity and/or correlation dimensions less than 5) that suggest deterministic chaos. These movement patterns appear regular, but at unpredictable times the patterns become disturbed before returning to regulation. There are both behavioral and physiological advantages to movements that follow a model of deterministic chaos control.     

Visualization of body thermoregulation by infrared imaging

This paper analyzes two mechanisms applied on the human body in order to study the thermoregulatory system according to heat generation and heat loss. Two approaches are presented. The first approach is based on plethysmography, where an armband is placed on the forearm in order to modulate the blood flow. The second approach uses a cold stimulation. The visualization is achieved using infrared imaging devices. The resulting images reveal a temperature balance between the stimulated and the non-stimulated hands. The thermal behavior and typical thermographic recordings on each subject are discussed and analyzed in response to different stimulations.     

Food consumption does not affect the preferred body temperature of Yarrow's spiny lizard (Sceloporus jarrovi)

When animals consume less food, they must reduce their body temperature to maximize growth. However, high temperatures enhance locomotion and other performances that determine survival and reproduction. Therefore, thermoregulatory behaviors during different metabolic states reveal the relative importance of conserving energy and sustaining performance. Using artificial thermal gradients, we measured preferred body temperatures of male spiny lizards (Sceloporus jarrovi) in fed and fasted states. Both the mean and maximal body temperatures (33° and 35 °C, respectively) were unaffected by metabolic state. This finding suggests that the benefits of foraging effectively, evading predators, and defending territory outweigh the energetic cost of a high body temperature during fasting.     

Body temperature and thermoregulation in two species of yellowjackets,Vespula germanica and V. maculifrons

In spite of the abundance and broad distribution of social wasps, little information exists concerning thermoregulation by individuals. We measured body temperatures of the yellowjackets Vespula germanica and V. maculifrons and examined their thermoregulatory mechanisms. V. germanica demonstrated thermoregulation via a decreasing gradient between thorax temperature and ambient temperature as ambient temperature increased. V. maculifrons exhibited a constant gradient at lower ambient temperatures but thorax temperature was constant at high ambient temperatures. Head temperature exhibited similar patterns in both species. In spite of low thermal conductances, a simple heat budget model predicts substantial heat loads in warm conditions in the absence of thermoregulation. Both species regurgitated when heated on the head. A smaller volume of regurgitant was produced at lower head temperatures and a larger volume at higher head temperatures. Small regurgitations resulted in stabilization of head temperature, while large ones resulted in 4°C decreases in head temperature. Regurgitation was rare when wasps were heated upon the thorax. Abdomen temperature was 3–4°C above ambient temperature, and approached ambient temperature under the hottest conditions. No evidence was found for shunting of hot hemolymph from thorax to abdomen as a cooling mechanism. The frequency of regurgitation in workers returning to the nest increased with ambient temperature. Regurgitation may be an important thermoregulatory strategy during heat stress, but is probably not the only mechanism used in yellowjackets.Abbreviations M _b body mass - M _th thorax mass - T _a ambient temperature - T _ab abdomen temperature - T _b body temperature - T _h head temperature - T _th thorax temperature - C _t thermal conductance     

Temperature selection by juvenile tuatara (Sphenodon punctatus) is not influenced by temperatures experienced as embryos

Most reptiles thermoregulate to achieve body temperatures needed for biological processes, such as digestion and growth. Temperatures experienced during embryogenesis may also influence post-hatching growth rate, potentially through influencing post-hatching choice of temperatures. We investigated in laboratory settings whether embryonic temperatures (constant 18 °C, 21 °C and 22 °C) influence selected body temperatures (T_sel) of juvenile tuatara (Sphenodon punctatus), providing a possible mechanism for differences in growth rates. We found that incubation temperature does not influence T_sel. Although the average daily mean T_sel was 21.6 ± 0.3 °C, we recorded individual T_sel values up to 33.5 °C in juvenile tuatara, which is higher than expected and above the panting threshold of 31–33 °C reported for adults. We found diel patterns of T_sel of juvenile tuatara, observing a general pattern of two apparent peaks and troughs per day, with T_sel being significantly lower around dawn and at 1500 h than any other time. When comparing our results with other studies on tuatara there is a remarkable consistency in mean T_sel of ~ 21 °C across tuatara of different ages, sizes and acclimatization histories. The ability of juvenile tuatara to withstand a wide range of temperatures supports their former widespread distribution throughout New Zealand and warrants further investigation into their plasticity to withstand climate warming, particularly where they have choices of habitat and the ability to thermoregulate.     

Geographic variation and acclimation effects on thermoregulation behavior in the widespread lizard Liolaemus pictus

Populations at the warm range margins of the species distribution may be at the greatest risks of extinction from global warming unless they can tolerate extreme environmental conditions. Yet, some studies suggest that the thermal behavior of some lizard species is evolutionarily rigid. During two successive years, we compared the thermal biology of two populations of Liolaemus pictus living at the northern (warmer) and one population living at the southern (colder) range limits, thus spanning an 800 km latitudinal distance. Populations at the two range margins belong to two deeply divergent evolutionary clades. We quantified field body temperatures (T_b), laboratory preferred body temperatures (PBT), and used operative temperature data (T_e) to calculate the effectiveness of thermoregulation (E). During one year in all populations, we further exposed half of the lizards to a cold or a hot acclimation treatment to test for plasticity in the thermal behavior. The environment at the southern range limit was characterized by cooler weather and lower T_e. Despite that, females had higher T_b and both males and females had higher PBT in the southernmost population (or clade) than in the northernmost populations. Acclimation to cold conditions led to higher PBT in all populations suggesting that plastic responses to thermal conditions, instead of evolutionary history, may contribute to geographic variation. Lizards regulated moderately well their body temperature (E≈0.7): they avoided warm microhabitats in the northern range but capitalized on warm microhabitats in the southern range. We review literature data to show that Liolaemus species increase their thermoregulation efficiency in thermally challenging environments. Altogether, this indicates that habitats of low thermal quality generally select against thermoconformity in these lizards.     

Behavioural thermoregulation and the relative roles of convection and radiation in a basking butterfly

Poikilothermic animals are often reliant on behavioural thermoregulation to elevate core-body temperature above the temperature of their surroundings. Butterflies are able to do this by altering body posture and location while basking, however the specific mechanisms that achieve such regulation vary among species. The role of the wings has been particularly difficult to describe, with uncertainty surrounding whether they are positioned to reduce convective heat loss or to maximise heat gained through radiation. Characterisation of the extent to which these processes affect core-body temperature will provide insights into the way in which a species׳ thermal sensitivity and morphological traits have evolved. We conducted field and laboratory measurements to assess how basking posture affects the core-body temperature of an Australian butterfly, the common brown (Heteronympha merope). We show that, with wings held open, heat lost through convection is reduced while heat gained through radiation is simultaneously maximised. These responses have been incorporated into a biophysical model that accurately predicts the core-body temperature of basking specimens in the field, providing a powerful tool to explore how climate constrains the distribution and abundance of basking butterflies.     

The peak of thermoregulation effectiveness: Thermal biology of the Pyrenean rock lizard,Iberolacerta bonnali (Squamata,Lacertidae)

We studied, at 2200 m altitude, the thermal biology of the Pyrenean rock lizard, Iberolacerta bonnali, in the glacial cirque of Cotatuero (National Park of Ordesa, Huesca, Spain). The preferred thermal range (PTR) of I. bonnali indicates that it is a cold-adapted ectotherm with a narrow PTR (29.20–32.77 °C). However, its PTR (3.57 °C) is twice as wide as other Iberolacerta lizards, which may be explained by its broader historical distribution. The studied area is formed by a mosaic of microhabitats which offer different operative temperatures, so that lizards have, throughout their entire daily period of activity, the opportunity to choose the most thermally suitable substrates. I. bonnali achieves an effectiveness of thermoregulation of 0.95, which makes it the highest value found to date among the Lacertidae, and one of the highest among lizards. Their relatively wide distribution, their wider PTR, and their excellent ability of thermoregulation, would make I. bonnali lizards less vulnerable to climate change than other species of Iberolacerta. Thanks to its difficult access, the studied area is not visited by a large number of tourists, as are other areas of the National Park. Thus, it is a key area for the conservation of the Pyrenean rock lizard. By shuttling between suitable microhabitats, lizards achieve suitable body temperatures during all day. However, such thermally suitable microhabitats should vary in other traits than thermal quality, such as prey availability or predation risk. Hence, it seems that these not-thermal traits are not constraining habitat selection and thermoregulation in this population. Therefore, future research in this population may study the causes that would lead lizards to prioritize thermoregulation to such extent in this population.     

Behavioural thermoregulation by subyearling fall (autumn) Chinook salmon Oncorhynchus tshawytscha in a reservoir

This study investigated behavioural thermoregulation by subyearling fall (autumn) Chinook salmon Oncorhynchus tshawytscha in a reservoir on the Snake River, Washington, U.S.A. During the summer, temperatures in the reservoir varied from 23° C on the surface to 11° C at 14 m depth. Subyearlings implanted with temperature-sensing radio transmitters were released at the surface at temperatures >20° C during three blocks of time in summer 2004. Vertical profiles were taken to measure temperature and depth use as the fish moved downstream over an average of 5·6–7·2 h and 6·0–13·8 km. The majority of the subyearlings maintained average body temperatures that differed from average vertical profile temperatures during most of the time they were tracked. The mean proportion of the time subyearlings tracked within the 16–20° C temperature range was larger than the proportion of time this range was available, which confirmed temperature selection opposed to random use. The subyearlings selected a depth and temperature combination that allowed them to increase their exposure to temperatures of 16–20° C when temperatures <16 and >20° C were available at lower and higher positions in the water column. A portion of the subyearlings that selected a temperature c. 17·0° C during the day, moved into warmer water at night coincident with an increase in downstream movement rate. Though subyearlings used temperatures outside of the 16–20° C range part of the time, behavioural thermoregulation probably reduced the effects of intermittent exposure to suboptimal temperatures. By doing so, it might enhance growth opportunity and life-history diversity in the population of subyearlings studied.     

Thermoregulatory behaviour in two species of iguanid lizards (Crotaphytus collaris andSauromalus obesus): diel variation and the effect of pinealectomy

Summary Thermal selection by collared lizards (Crotaphytus collaris) and chuckawallas (Sauromalus obesus) was studied in continuously operating laboratory thermal gradients in a 12L12D photoperiod in spring-summer. In both species, selected temperatures above the median were higher during the day than at night. Below the median selected temperature, however, nighttime temperature significantly exceeded daytime temperature in chuckawallas but not in collared lizards. Pinealectomized collared lizards selected significantly lower temperatures than sham-operated or intact controls. This effect was most pronounced at night. Pinealectomy had no effect on the temperatures selected by chuckawallas. The significance of these results is discussed in relation to the seasonal life-history strategy of these two species.