Testing the fitness consequences of the thermoregulatory and parental care models for the origin of endothermy

The origin of endothermy is a puzzling phenomenon in the evolution of vertebrates. To address this issue several explicative models have been proposed. The main models proposed for the origin of endothermy are the aerobic capacity, the thermoregulatory and the parental care models. Our main proposal is that to compare the alternative models, a critical aspect is to determine how strongly natural selection was influenced by body temperature, and basal and maximum metabolic rates during the evolution of endothermy. We evaluate these relationships in the context of three main hypotheses aimed at explaining the evolution of endothermy, namely the parental care hypothesis and two hypotheses related to the thermoregulatory model (thermogenic capacity and higher body temperature models). We used data on basal and maximum metabolic rates and body temperature from 17 rodent populations, and used intrinsic population growth rate (R(max)) as a global proxy of fitness. We found greater support for the thermogenic capacity model of the thermoregulatory model. In other words, greater thermogenic capacity is associated with increased fitness in rodent populations. To our knowledge, this is the first test of the fitness consequences of the thermoregulatory and parental care models for the origin of endothermy.     

Endothermy in African platypleurine cicadas: the influence of body size and habitat (Hemiptera: Cicadidae)

The platypleurine cicadas have a wide distribution across Africa and southern Asia. We investigate endothermy as a thermoregulatory strategy in 11 South African species from five genera, with comparisons to the lone ectothermic platypleurine we found, in an attempt to ascertain any influence that habitat and/or body size have on the expression of endothermy in the platypleurine cicadas. Field measurements of body temperature (T(b)) show that these animals regulate T(b) through endogenous heat production. Heat production in the laboratory elevated T(b) to the same range as in animals active in the field. Maximum T(b) measured during calling activity when there was no access to solar radiation ranged from 13.2 degrees to 22.3 degrees C above ambient temperature in the five species measured. The mean T(b) during activity without access to solar radiation did not differ from the mean T(b) during diurnal activity. All platypleurines exhibit a unique behavior for cicadas while warming endogenously, a temperature-dependent telescoping pulsation of the abdomen that probably functions in ventilation. Platypleurines generally call from trunks and branches within the canopy and appear to rely on endothermy even when the sun is available to elevate T(b), in contrast to the facultative endothermy exhibited by New World endothermic species. The two exceptions to this generalization we found within the platypleurines are Platypleura wahlbergi and Albanycada albigera, which were the smallest species studied. The small size of P. wahlbergi appears to have altered their thermoregulatory strategy to one of facultative endothermy, whereby they use the sun when it is available to facilitate increases in T(b). Albanycada albigera is the only ectothermic platypleurine we found. The habitat and host plant association of A. albigera appear to have influenced the choice of ectothermy as a thermoregulatory strategy, as the species possesses the metabolic machinery to elevate to the T(b) range observed in the endothermic species. Therefore, size and habitat appear to influence the expression of thermoregulatory strategies in African platypleurine cicadas.     

Thermoregulatory responses of two mouse Mus musculus strains selectively bred for high and low food intake

We examined the thermoregulatory responses of male and female mice Mus musculus that had been divergently selected on voluntary food intake, corrected for body mass, to produce a high-intake and a low-intake strain. Resting metabolic rate was determined by indirect calorimetry (at 30 degrees C, 25 degrees C, 15 degrees C and 5 degrees C). Body temperature responses were measured in a separate group of mice in a parallel protocol. High-intake mice had significantly elevated body masses compared to low-intake mice in both sexes. Lower critical temperature in both strains appeared to be around 28 degrees C. At 30 degrees C there was a significant strain effect on resting metabolic rate, with high strain mice having greater metabolism than low strain mice. Sex and body mass were not significant main effects on resting metabolic rate and there were no significant interactions. Body temperature measured at 30 degrees C, 25 degrees C, 15 degrees C and 5 degrees C differed significantly between sexes (females higher) and there was a significant sexxbody mass interaction effect, but there was no difference between strains. Thermal conductance was significantly related to strain and sex, mice from the high strain and males having greater thermal conductances than mice from the low strain and females. Artificial selection has resulted in high-intake mice having greater body masses and greater thermal conductances, which together account for up to 45% of the elevated daily energy demands that underpin the increase in food intake. The greater levels of food intake were also associated with higher resting metabolic rates at 30 degrees C.     

Thermoregulation of weaned northern elephant seal (Mirounga angustirostris) pups in air and water

Fasting weaned northern elephant seal pups (Mirounga angustirostris) experience diverse environmental conditions on land and in water on a daily basis. Each environment undoubtedly induces distinct energetic costs that may vary for pups of differing body condition. To determine the energetic costs associated with different environmental conditions and whether costs vary between individuals, body mass, surface area, volume, body composition, resting metabolic rate, and core body temperature were determined for 17 weaned northern elephant seal pups from A?o Nuevo, California. Metabolic rate and body temperature were measured for pups resting in air (20.9 degrees +/-0.8 degrees C), cold water (3.8 degrees+/-0.4 degrees ;C), and warm water (14.5 degrees+/-0.2 degrees C). Resting metabolic rate increased with body mass (range: 62.0-108.0 kg) and was also correlated with lean mass and lipid mass. Metabolic rates ranged from 293.6 to 512.7 mL O(2) min(-1) and were lowest for pups resting in cold water. Thermal conductance, calculated from metabolic rate and core body temperature, ranged from 3.1 to 15.2 W degrees C(-1), with the highest values in air and the lowest values in cold water. Metabolic responses to the three environmental conditions did not differ with individual variation in body condition. For all elephant seal pups, a consequence of high lipid content is that thermoregulatory costs are greatest on land and lowest in cold water, a pattern that contrasts markedly with terrestrial mammals.     

THE EVOLUTION OF ENDOTHERMY: TESTING THE AEROBIC CAPACITY MODEL

One of the most important events in vertebrate evolution was the acquisition of endothermy, the ability to use metabolic heat production to elevate body temperature above environmental temperature. Several verbal models have been proposed to explain the selective factors leading to the evolution of endothermy. Of these, the aerobic capacity model has received the most attention in recent years. The aerobic capacity model postulates that selection acted mainly to increase maximal aerobic capacity (or associated behavioral abilities) and that elevated resting metabolic rate evolved as a correlated response. Here we evaluate the implicit evolutionary and genetic assumptions of the aerobic capacity model. In light of this evaluation, we assess the utility of phenotypic and genetic correlations for testing the aerobic capacity model. Collectively, the available intraspecific data for terrestrial vertebrates support the notion of a positive phenotypic correlation between resting and maximal rates of oxygen consumption within species. Interspecific analyses provide mixed support for this phenotypic correlation. We argue, however, that assessments of phenotypic or genetic correlations within species and evolutionary correlations among species (from comparative data) are of limited utility, because they may not be able to distinguish between the aerobic capacity model and plausible alternatives, such as selection acting directly on aspects of thermoregulatory abilities. We suggest six sources of information that may help shed light on the selective factors important during the evolution of high aerobic metabolic rates and, ultimately, the attainment of endothermy. Of particular interest will be attempts to determine, using a combination of mechanistic physiological and quantitative-genetic approaches, whether a positive genetic correlation between resting and maximal rates of oxygen consumption is an ineluctable feature of vertebrate physiology.     

Antipyresis due to centrally administered vasopressin differentially alters thermoregulatory effectors depending on the ambient temperature

The intracerebroventricular (i.c.v.) administration of arginine vasopressin (AVP), in the febrile rat elicits an antipyresis at cold, warm and neutral ambient temperatures. These experiments were conducted, therefore, to elucidate the thermoregulatory effector mechanisms responsible for this antipyretic effect. At 25 degrees C, AVP-induced antipyresis was mediated by tail skin vasodilation while metabolic rate was unaffected. At 4 degrees C, the antipyresis produced by AVP was approximately double that seen at 25 degrees C. This effect appeared to be mediated exclusively by inhibition of heat production since the metabolic rate decreased markedly following AVP. This antipyresis at 4 degrees C was accompanied by cutaneous vasoconstriction. At 32 degrees C, neither vasomotor tone, metabolic rate nor evaporative heat loss could be shown to contribute to the small antipyretic effect elicited by AVP. We conclude from these data that i.c.v. AVP is producing antipyresis by affecting the febrile body temperature set-point mechanism since the thermoregulatory strategy to lose heat varies at different ambient temperatures and the decrease in body temperature cannot be shown to be due to changes in a single effector mechanism.     

The origin of homoiothermy--unsolved problem

The analysis of allometric dependence of energy expenditure on body mass among reptiles, birds and mammals has shown that standard metabolic rate of reptiles when they are warmed up to the temperature of homoiothermic animals is an order of magnitude lower than that of birds and mammals. Basal metabolism is originated as special feature historically related to the metabolism during active behavior, rather than thermal regulation. Facultative endothermy was not advantageous for large animals because of long time needed to warm up the body. The ancestors of birds and animals escaped negative consequences of van't-Hoff equation by choosing constant body temperature. Heat conductivity of reptile's covers is so great, that it cannot keep endogenous warm of resting animal at any temperature of the body. Reptile "dressed" in covers of bird or mammal would be able to keep warm under conditions of maximal aerobic muscular activity and body temperature similar to that of homoiothermic animals. The base of chemical thermoregulation in birds and mammals is a thermoregulatory muscle tonus which remains unknown. One can suppose that during evolution of birds and mammals the saltation-liked origin of endothermy "fixed" the level of metabolism typical for running reptile and transformed in into the basal metabolism. This event took place at the cell and tissue level. The absence of palaeontological evidences and intermediate forms among recent species does not allow easy understanding of homoiothermy origin.     

Developmental and adult acclimation effects of ambient temperature on temperature regulation of mice selected for high and low levels of nest-building

Summary Genetic and environmental components of adaptation to cold inMus musculus were assessed in a study of the effects of selective breeding for behavioral temperature regulation (indexed by high and low levels of nest-building), rearing mice from birth in the cold, and cold acclimation of adult animals, on thermoregulatory traits. Mice from the eleventh selected generation of a high-nesting line maintained higher resting metabolic rates and body temperatures, while at the same time consuming less food when compared with mice from the low-nesting line (Table 1). High-nesting mice were also more discriminating in their temperature preference when placed on a thermal gradient. Thus, common genetic loci must influence a variety of energy conservation measures important for survival in the cold, including insulative nest-building, metabolic efficiency, and optimum microhabitat selection.Rearing mice at 5°C from birth until 70 days of age resulted in permanent increases in nonshivering thermogenesis, weight of interscapular brown adipose tissue, and core body temperature when compared to mice raised at 22°C (Table 1). These greater heat production capacities were accompanied by consumption of more food. Cold acclimation of adults at 5°C for 3 weeks similarly increased measures of thermogenic capacity (nonshivering thermogenesis and interscapular brown adipose tissue) as well as food consumption, when compared to the effects of warm acclimation, but differed from the effects of cold-rearing in that while resting metabolic rates were elevated, no significant differences in body temperature were found (Table 1).Sex differences were also noted for most of the thermoregulatory measures, with the lighter females scoring higher on thermal preference, resting metabolic rate, nonshivering thermogenesis, brown fat, and food consumption.In general, these results suggest that a more precise partitioning of the genetic and environmental factors which influence thermoregulatory traits in mammals could eventually result in a better understanding of the differences which exist between acclimated and acclimatized animals.     

动物内温性进化研究进展

对动物内温性进化的研究进行了较为系统的论述,包括内温性动物概念的由来、特点和起源的选择因子。内温性起源的选择因子包括8个模型：热生态位扩展模型、恒温与代谢效率模型、降低个体大小模型、姿势改变模型、增加脑大小模型、有氧呼吸能力模型、双亲行为模型和同化能力模型。其中后3个模型较为重要。有氧呼吸能力模型认为,选择提高支持物理运动的最大呼吸能力,而增加的静止代谢作为其相关反应而得以进化。该假说得到种内研究数据的支持,而种问的数据并小完全支持。双亲行为模型是指在鸟兽类中,内温性是对双亲行为选择的结果,因为内温性为双亲控制抚育温度提供了保证。同化能力模型认为,在鸟类和兽类中内温性进化由以下两个因素所推动：①子代出生后双亲行为加强;②为支持每日总体能量高速消耗所需,动物内脏器官能力增强而导致的较高维持消耗。     

Prediction of the thermoregulatory response for clothed immersion in cold water

A multi-compartmental thermoregulatory model was applied to data of ten resting clothed males immersed for 3 h in water at 10 and 15 degrees C. Clothing consisted of a dry suit and either a light or heavy undergarment, representing a total insulation of 0.15 (0.95) or 0.20 m2 degrees CW-1 (1.28 clo), respectively. Data were grouped according to low (less than 14%) and high (14 to 24%) body fat individuals. Mean decreases in rectal temperature ranged from 0.79 to 1.38 degrees C, mean decreases in the mean weighted skin temperature ranged from 6.3 to 10.2 degrees C, and mean increases in the metabolic rate ranged from 33.9 to 80.8 W. The model consists of eight segments, each representing a specific region of the body. Each segment is comprised of compartments representing the core, muscle, fat, skin, and clothing. Each compartment is assigned thermophysical values of heat conduction and heat capacitance, and with the exception of clothing, physiological values of blood flow and metabolic heat production. During cold exposure, responses are directed towards increased heat production in the form of shivering and heat conservation in the form of vasoconstriction and convective heat exchange at the vascular level. Agreement between the model predictions and the experimental observations was obtained by adjusting the parameters governing these responses.(ABSTRACT TRUNCATED AT 250 WORDS)     

Metabolic responses of hibernating golden-mantled ground squirrels Citellus lateralis to lowered environmental temperatures

1. Hibernating C. lateralis were exposed to lowered ambient temperatures in order to investigate the relationship between hibernation stress and the thermoregulatory responses of the animals. 2. The least hibernation-stress squirrels exhibited a passive decline in metabolic rate until their body temperatures stabilized close to microenvironmental temperature. 3. The most stressed individuals aroused from hibernation in response to the declining ambient temperatures. 4. Intermediately stressed animals demonstrated an initial passive decline in temperature; however, at various temperatures (0.3-6.8 degrees C), this group increased their metabolic rate but did not arouse from hibernation.     

Metabolic depression and heat balance in starving Wistar rats

Resting metabolic rate and heat balance was studied in rats starved for 8 days at ambient temperature 22 degrees C and 30 degrees C. A depression of the resting metabolic rate was observed, at both temperatures. Metabolic rate depression, expressed as a function of the ratio between the real body wt and the normal body wt, was less at 22 degrees C than at 30 degrees C. Deep body temperature decrements of 2 degrees C and 0.6 degrees C by the end of starvation indicated that central temperature controlling mechanisms were affected. Concurrent decrements of evaporative heat loss did not account for the changes in heat conductance, thus indicating that a reduction of peripheral blood circulation also took part.     

Temperature,metabolic power and the evolution of endothermy

Endothermy has evolved at least twice, in the precursors to modern mammals and birds. The most widely accepted explanation for the evolution of endothermy has been selection for enhanced aerobic capacity. We review this hypothesis in the light of advances in our understanding of ATP generation by mitochondria and muscle performance. Together with the development of isotope‐based techniques for the measurement of metabolic rate in free‐ranging vertebrates these have confirmed the importance of aerobic scope in the evolution of endothermy: absolute aerobic scope, ATP generation by mitochondria and muscle power output are all strongly temperature‐dependent, indicating that there would have been significant improvement in whole‐organism locomotor ability with a warmer body. New data on mitochondrial ATP generation and proton leak suggest that the thermal physiology of mitochondria may differ between organisms of contrasting ecology and thermal flexibility. Together with recent biophysical modelling, this strengthens the long‐held view that endothermy originated in smaller, active eurythermal ectotherms living in a cool but variable thermal environment. We propose that rather than being a secondary consequence of the evolution of an enhanced aerobic scope, a warmer body was the means by which that enhanced aerobic scope was achieved. This modified hypothesis requires that the rise in metabolic rate and the insulation necessary to retain metabolic heat arose early in the lineages leading to birds and mammals. Large dinosaurs were warm, but were not endotherms, and the metabolic status of pterosaurs remains unresolved.     

Metabolism and thermoregulation in the Cabrera vole (Rodentia: Microtus cabrerae)

Metabolism and thermoregulation were studied for the first time in the Cabrera vole (Microtus cabrerae), an endemic and threatened rodent of the Iberian Peninsula. Low values of resting metabolic rate (RMR) were registered (1.13 mlO(2) g(-1) h(-1)) at the lower limit of the thermoneutral zone (TNZ) (around 33.5 degrees C). Body temperature increased near the TNZ up to 37.3 degrees C but remained stable, around 36 degrees C, at ambient temperatures below 25 degrees C. Values of thermal conductance remained quite stable at ambient temperatures of 10-25 degrees C (0.144-0.160 mlO(2) g(-1) h(-1) degrees C) and increased to 0.301 mlO(2) g(-1) h(-1) degrees C at 33.5 degrees C. Data revealed that M. cabrerae developed a highly adaptive ability of conserving energy and lowering the metabolic cost of thermoregulation at high ambient temperatures, allowing the body temperature to approximate that of the environment and exhibiting low resting metabolic rate and high conductance.     

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]。     

Thermoregulatory and nonthermoregulatory heat production in burned rat

Severely burned patients are hypermetabolic within their thermoneutral zone (TNZ), where there are no thermoregulatory demands on heat production. The rat has been used as a model of postburn hypermetabolism without clear evidence that it behaves in a similar way. Male rats (400-500 g; n = 34-39) were placed as a group in a respiration chamber and metabolic rates for the average rat were determined over 3-6 h at ambient temperatures between 9 and 36 degrees C. Colonic temperatures (Tco) and body weights were measured after each run. Animals were studied sequentially as normals (N), after clipping (C) and following 50% total body surface scald burns. Clipping increased the lower critical temperature (LCT) from 27.7 to 29.1 degrees C without affecting resting heat production (N = 42.6 +/- 0.5; C = 42.0 +/- 0.8 W/m2; mean +/- S.E.) or Tco (N = 36.6 +/- 0.1; C = 36.6 +/- 0.1 degrees C) in the TNZ. Injury increased LCT to 32.8 degrees C and the burned animals were hypermetabolic (47.2 +/- 0.6 W/m2; P less than 0.05 vs. N) and febrile (36.9 +/- 0.1 degrees C; P less than 0.05 vs. N) in the elevated TNZ. These metabolic and temperature responses of burned rats are limited in magnitude but are qualitatively similar to those of patients. The extra heat production in the TNZ reflects the basic metabolic cost of injury.     

Oscillatory pattern in oxygen consumption of Hummingbirds

Mammals and birds offer the most conspicuous example of homeothermic endothermy, a metabolic feature that implies maintenance of a constant body temperature along broad ranges of ambient temperature. The concept of homeothermic endothermy has been developed in close association with the terms thermoneutral zone and basal metabolic rate. These two metabolic parameters, however, are not easily estimated in micro-endotherms, a difficulty that might emerge from intrinsic aspects of endothermy in minute animals. To address this issue, we used empirical work derived from theoretical considerations. Our theoretical analysis is based on a model of body temperature control by shifts in metabolic rate, and assumes that micro-endotherms lose heat very quickly due to body size, and exhibit a remarkable capacity to rapidly increase metabolic output. We found that these two metabolic traits can lead to non-equilibrium metabolic rate and body temperature. We then measured metabolic rate and body temperature during euthermia in two species of hummingbirds, and analyzed data using the χ² periodogram statistic and a power spectral analysis. We found long-range correlation in both oxygen consumption and body temperature during euthermia, a finding that suggests non-random 1/f oscillations. A similar pattern was not found in the rat, a much larger endotherm. Hummingbirds, then, do not appear to maintain steady-state metabolic conditions during euthermia. If, as we suggest, this pattern applies to micro-endotherms in general, the traditional concepts of thermoneutral zone and basal rate of metabolism might not apply to these animals.     

Thermoregulatory consequences of long-term microwave exposure at controlled ambient temperatures

This study was designed to identify and measure changes in thermoregulatory responses, both behavioral and physiological, that may occur when squirrel monkeys are exposed to 2450-MHz continuous wave microwaves 40 hr/week for 15 weeks. Power densities of 1 or 5 mW/cm2 (specific absorption rate = 0.16 W/kg per mW/cm2) were presented at controlled environmental temperatures of 25, 30, or 35 degrees C. Standardized tests, conducted periodically, before, during, and after treatment, assessed changes in thermoregulatory responses. Dependent variables that were measured included body mass, certain blood properties, metabolic heat production, sweating, skin temperatures, deep body temperature, and behavioral responses by which the monkeys selected a preferred environmental temperature. Results showed no reliable alteration of metabolic rate, internal body temperature, blood indices, or thermoregulatory behavior by microwave exposure, although the ambient temperature prevailing during chronic exposure could exert an effect. An increase in sweating rate occurred in the 35 degrees C environment, but sweating was not reliably enhanced by microwave exposure. Skin temperature, reflecting vasomotor state, was reliably influenced by both ambient temperature and microwaves. The most robust consequence of microwave exposure was a reduction in body mass, which appeared to be a function of microwave power density.     

Simple roost nests confer large energetic savings for sparrow-weavers

White-browed sparrow-weavers (Plocepasser mahali, body mass 40 g) are group-living passerines adapted to the semi-arid environment of north-eastern and south-western Africa. During winter, the nocturnal ambient temperature of these regions often falls below 0 degrees C. imposing conditions demanding an increase in thermoregulatory heat production. Individuals roost throughout the year in inverted U-shaped roost nests. We investigated the energetic advantages of roosting by measuring nest and ambient temperatures in the field, as well as the resting metabolic rate (RMR) of the birds. The sparrow-weavers exhibited a wide thermoneutral zone (13 degrees C - 32 degrees C). Although RMR at thermoneutrality (40.2 J g.h(-1)) conforms with those of other passerines. the value at 0 degrees C (74.8 J g.h(-1)) is significantly lower than expected. The slope of the line below the lower critical temperature is unexpectedly steep, however, and appears to cause the physiological requirement for nest roosting due to a high cost of thermoregulation at low temperatures, perhaps due to shivering or non-shivering thermogenesis. The nest temperature at 0 degrees C ambient is 5 degrees C. resulting in a saving of some 7% in the energy spent during winter nights when food resources are in short supply compared with the rest of the year.     

Shifts in thermoregulatory patterns with pregnancy in the poikilothermic mammal—The naked mole-rat (Heterocephalus glaber)

1. 1. The naked mole-rat (Heterocephalus glaber) is a poikilothermic mammal. During gestation metabolic shifts that differ from both mammalian and reptilian thermoregulatory patterns occurred.

2. 2. Body temperature was directly dependent on ambient temperature. At low ambient temperatures the temperature differential (T_b − T_a) was approximately 3°C, whereas at higher ambient temperatures the temperature differential diminished.

3. 3. In early pregnancy (prior to week 3) oxygen consumption at low ambient temperatures was greater than that of non-reproductive animals. A maximal metabolic rate (3.2 ± 1.0 ml O₂ . g⁻¹ . h⁻¹) occurred at an ambient temperature of 27°C. Thereafter the endothermic pattern of metabolism with increasing ambient temperatures was evident. Oxygen consumption decreased with increasing ambient temperature to minimal rates of 1.2 ± 0.1 ml O₂ . g⁻¹ . h⁻¹ over the ambient temperature range of 31–34°C.

4. 4. Oxygen consumption in late pregnancy (1.8 ± 0.1 ml O₂ . g⁻¹ . h⁻¹) was not correlated with ambient temperature over the entire ambient temperature range measured (24–36°C).

5. 5. Differences in thermoregulation in early and late pregnancy may be attributed to different rates of heat loss as a consequence of (a) changes in surface area and body mass or (b) vascular changes. Furthermore the thermoregulatory changes in late pregnancy may indicate that maximal overall metabolic capacity had been reached, for peak resting metabolism (expressed per animal rather than per gram body mass) in early pregnancy was similar to observed metabolism in late pregnancy.

6. 6. The dissociation of metabolism from both ambient temperature and body temperature in late pregnancy could confer an energetic advantage to this arid dwelling underground inhabitant; for it may enable the breeding female to partition a greater portion of available energy into reproduction.