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High-altitude environments are cold and hypoxic, and many high-altitude natives have evolved changes in respiratory physiology that improve O2 uptake in hypoxia as adults. Altricial mammals undergo a dramatic metabolic transition from ectothermy to endothermy in early post-natal life, which may influence the ontogenetic development of respiratory traits at high altitude. We examined the developmental changes in respiratory and haematological traits in deer mice (Peromyscus maniculatus) native to high altitude, comparing the respiratory responses to progressive hypoxia between highland and lowland deer mice. Among adults, highlanders exhibited higher total ventilation and a more effective breathing pattern (relatively deeper tidal volumes), for mice that were caught and tested at their native altitudes and those lab-raised in normoxia. Lab-raised progeny of each population were also tested at post-natal day (P)7, 14, 21 and 30. Highlanders developed an enhanced hypoxic ventilatory response by P21, concurrent with the full maturation of the carotid bodies, and their more effective breathing pattern arose by P14; these ages correspond to critical benchmarks in the full development of homeothermy in highlanders. However, highlanders exhibited developmental delays in ventilatory sensitivity to hypoxia, hyperplasia of type I cells in the carotid body and increases in blood haemoglobin content compared with lowland mice. Nevertheless, highlanders maintained consistently higher arterial O2 saturation in hypoxia across development, in association with increases in blood–O2 affinity that were apparent from birth. We conclude that evolved changes in respiratory physiology in high-altitude deer mice become expressed in association with the post-natal development of endothermy.

Endothermy provides considerable benefits to an organism but requires large energy investment. To understand potential driving forces that would lead to the evolution of endothermy, it is important to understand the energy costs and potential benefits of intermediate steps between ectothermy and homeothermic endothermy as well as the influences of environmental conditions on energetic costs. However, efforts to examine intermediate conditions are greatly limited by the predominant natural dichotomy between ectothermy and endothermy. Facultative endothermy by brooding pythons provides a fortunate study system where endothermy is beneficial but not essential. As one cannot control the extent of energy investment in heat production by a female python, we created an artificial snake with controllable heating capability. This enabled us to determine the energetic costs of maintaining a clutch at a preferred temperature, and to determine the relative thermal benefit of limited energy-producing capability (i.e. 50% of the required energy to maintain the preferred developmental temperature). We manipulated the pseudoserpent's clutch size (5, 10, 15 eggs), diel ambient temperature cycle (2, 4, 6°C) and insulation (with and without) at each of these power levels: unlimited power, half required power and no power. We found no significant effect of clutch size on either power requirements or developmental temperature. Energy requirements increased with the amplitude of the diel cycle and decreased with the addition of insulation, while the quality of the thermal environment decreased with the amplitude of the diel cycle. Interestingly, the quality of the thermal environment also decreased with the addition of insulation. We discuss these results within the context of the reproductive model of the evolution of endothermy.

According to theoretical predictions, endothermic homeotherms can be classified as either thermal specialists or thermal generalists. In high cost environments, thermal specialists are supposed to be more prone to using facultative heterothermy than generalists. We tested this hypothesis at the intraspecific level using male laboratory mice (C57BL/cmdb) fasted under different thermal conditions (20 and 10°C) and for different time periods (12–48 h). We predicted that variability of body temperature ( T b ) and time spent with T b below normothermy would increase with the increase of environmental demands (duration of fasting and cold). To verify the above prediction, we measured T b and energy expenditure of fasted mice. We did not record torpor bouts but we found that variations in T b and time spent in hypothermia increased with environmental demands. In response to fasting, mice also decreased their energy expenditure. Moreover, animals that showed more precise thermoregulation when fed had more variable T b when fasted. We postulate that the prediction of the thermoregulatory generalist–specialist trade-off can be applied at the intraspecific level, offering a valid tool for identifying mechanistic explanations of the differences in animal responses to variations in energy supply.

Summary Exposure of homeothermic animals to low environmental temperature is associated with oxidative stress in several body tissues. Because cold exposure induces a condition of functional hyperthyroidism, the observation that tissue oxidative stress also happens in experimental hyperthyroidism, induced by 3,5,3′-triiodothyronine (T3) treatment, suggests that this hormone is responsible for the oxidative damage found in tissues from cold-exposed animals. Examination of T3-responsive tissues, such as brown adipose tissue (BAT) and liver, shows that changes in factors favoring oxidative modifications are similar in experimental and functional hyperthyroidism. However, differences are also apparent, likely due to the action of physiological regulators, such as noradrenaline and thyroxine, whose levels are different in cold-exposed and T3-treated animals. To date, there is evidence that biochemical changes underlying the thermogenic response to cold as well as those leading to oxidative stress require a synergism between T3- and noradrenaline-generated signals. Conversely, available results suggest that thyroxine (T4) supplies a direct contribution to cold-induced BAT oxidative damage, but contributes to the liver response only as a T3 precursor.

SUMMARY Virtually all aspects of insect biology are affected by body temperature,and many taxa have evolved sophisticated temperature-control mechanisms. All insects, however, begin life as eggs and lack the ability to thermoregulate. Eggs laid on leaves experience a thermal environment, and thus a body temperature, that is strongly influenced by the leaves themselves. Because plants can maintain leaf temperatures that differ from ambient, e.g. by evapotranspiration, plant hosts may protect eggs from extreme ambient temperatures. We examined the degree to which leaves buffer ambient thermal variation and whether that buffering benefits leaf-associated insect eggs. In particular, we: (1) measured temperature variation at oviposition sites in the field, (2) manipulated temperatures in the laboratory to determine the effect of different thermal conditions on embryo development time and survival, and(3) tested embryonic metabolic rates over increasing temperatures. Our results show that Datura wrightii leaves buffer Manduca sexta eggs from fatally high ambient temperatures in the southwestern USA. Moreover, small differences in temperature profiles among leaves can cause large variation in egg metabolic rate and development time. Specifically, large leaves were hotter than small leaves during the day, reaching temperatures that are stressfully high for eggs. This study provides the first mechanistic demonstration of how this type of leaf-constructed thermal refuge interacts with egg physiology.

We used miniature data loggers implanted in the abdominal cavity to measure core body temperatures at 30 min intervals in eight (three males, five females) adult free-ranging springbok Antidorcas marsupialis in their natural habitat, over a period of 11-13 months. The animals were subjected to a nychthemeral range of air temperature that often exceeded 20 degrees C, with an absolute minimum temperature of -6 degrees C and a maximum of 34 degrees C. Abdominal temperature exhibited a low amplitude (approximately 1.2 degrees C) nychthemeral rhythm, with a temperature peak near sunset and a trough shortly after sunrise. The amplitude of the nychthemeral rhythm of body temperature was not correlated with the 24 h range of air temperature. Although mean 24 h body temperatures were positively correlated with corresponding air temperatures, mean daily body temperature increased, on average, by only 0.02 degrees C per 1 degrees C increase in air temperature, so that it was only approximately 0.3 degrees C higher in summer than in winter. Mean monthly body temperatures were strongly positively correlated with photoperiod and, in parallel with changes in the time of sunrise, the times at which the minimum and maximum body temperatures occurred were shifted approximately 1.2 h earlier in summer than in winter. Annual and daily variations in body temperature of springbok, like those of other free-living African ungulates, therefore appear to reflect an endogenous rhythm, entrained by the light:dark cycle, but largely independent of fluctuations in the environmental thermal load. Springbok exhibit remarkable homeothermy and do not employ adaptive heterothermy to survive in their natural environment.

SUMMARY An adaptation strategy whereby O 2 partial pressure, P O 2 , in the tissues is maintained within a low, narrow range of 1–3 kPa, largely independent of the inspired P O 2 , has been reported in water- and air-breathing poikilotherms and in homeotherms. Based on the postulate that this basic cellular mechanism has been established since the early stages of evolution, it has been hypothesized that it could be the consequence of an early adaptation strategy to maintain cellular oxygenation within the same low and primitive range. To test this hypothesis we studied the basic mechanisms of oxygen regulation in podocopid ostracods, minute crustaceans that have existed on earth for at least 500 million years. Podocopids lack any regulatory mechanism for adapting their ventilation to cope with changes in water oxygenation, and instead adjust their tissue oxygenation status by migrating through the O 2 gradient to sediment layers where the P O 2 of the water is 3–5 kPa. Experimental manipulation of the O 2 profile induced their vertical migration to follow this precise water P O 2 and demonstrates the existence of a regulation strategy. This strategy must be associated with the lower P O 2 values within the animal9s carapace valves, showing that podocopids can actively regulate their tissue P O 2 at constant but even lower values than the water. In conclusion, the low tissue P O 2 strategy could have existed in early crustaceans and, by extension, in early animals.

Homeothermic mammals experience pulmonary surfactant dysfunction with relatively small fluctuations in body temperature. However, ground squirrels survive dramatic changes in body temperature during hibernation, when body temperature drops from 37 degrees C to 0-5 degrees C during prolonged torpor bouts. Using type II cells isolated from both warm-active and torpid squirrels, we determined the effect of assay temperature, autonomic agonists and torpor on surfactant secretion. Basal secretion was significantly higher in type II cells isolated from torpid squirrels compared with warm-active squirrels when assayed at the body temperature of the animal from which they were isolated (4 degrees C and 37 degrees C, respectively). A change in assay temperature significantly decreased surfactant secretion. However, the change in secretory rate between 37 degrees C and 4 degrees C was less than expected if due to temperature alone (Q(10) range=0.8-1.2). Therefore, the surfactant secretory pathway in squirrel type II cells demonstrates some temperature insensitivity. When incubated at the body temperature of the animal from which the cells were isolated, the adrenergic agonist, isoproterenol, significantly increased surfactant secretion in both warm-active and torpid squirrel type II cells. However, the cholinergic agonist, carbamylcholine chloride, only increased secretion in torpid squirrel type II cells when incubated at 4 degrees C. Torpor did not affect basal cAMP production from isolated type II cells. However, the production of cAMP appears to be upregulated in response to isoproterenol in torpid squirrel type II cells. Thus, at the cellular level, both the secretory and regulatory pathways involved in surfactant secretion are thermally insensitive. Upregulating basal secretion and increasing the sensitivity of type II cells to cholinergic stimulation may be adaptative characteristics of torpor that enable type II cells to function effectively at 0-5 degrees C.

Homeothermic endothermy, the maintenance of a high and stable body temperature (Tb) using heat produced by elevated metabolism, is energetically expensive. There is increasing evidence that the earliest endotherms were heterotherms that, rather than maintaining strict homeothermy, allowed Tb to fluctuate with large variations between active and rest-phase Tb. The high level of homeothermy observed in modern mammals is therefore likely to have evolved from an ancestral heterothermic state. One of the hypotheses for the evolution of endothermy is that homeothermy allows for greater energetic output during reproduction (Parental Care Model). We tested this hypothesis by measuring metabolic rates over a range of ambient temperatures in both reproductive and non-reproductive Greater hedgehog tenrecs (Setifer setosus), a physiologically primitive mammal from Madagascar. Tenrecs have some of the lowest metabolic rates and highest levels of Tb variability of any mammal and are therefore good models of the ancestral eutherian state. During pregnancy and lactation there was an increase in metabolism and Tb below the thermoneutral zone, accompanied by a decrease in Tb variability. The lower critical limit of the thermoneutral zone was estimated at ~25°C. However, whereas increases in resting metabolism were substantial below 20°C (up to 150% higher during reproduction), daytime rest-phase ambient temperatures at the study site rarely reached equivalent low levels. Thus, S. setosus provide an example for how relatively low-cost increases in homeothermy could have led to substantial increases in fitness by allowing for the faster production of young. The mechanisms necessary for increases in thermogenesis during reproduction would have further benefited the development of homeothermy in mammals.

From an evolutionary perspective, the high basal metabolic rate (BMR) of homeotherms is hypothesised to be a by-product of natural selection for effective parental care. We estimated the daily milk output during two consecutive lactation bouts in mice divergently selected for high/low BMR and applied a cross-fostered design to control for potential differences in the between-line suckling abilities of nursed juveniles. Additionally, to remedy the potential limitation imposed by the ability of mother mice to dissipate excess heat, we exposed them to an ambient temperature of 17°C during the most energetically demanding second week of lactation. We found that the mice selected for high BMR produced significantly more milk in a 24 hour period in both reproductive bouts. The milk samples obtained from the high BMR females had lower protein concentration and did not differ with respect to fat. On the other hand, the concentration of the primary milk carbohydrate - lactose, was higher. Although all the above between-line differences were statistically significant, their magnitude was too small to unambiguously ascribe them as stemming from a positive genetic correlation between the physiological traits underlying BMR and lactation performance. Nevertheless, our study lends such support at least at the level of phenotypic variation.

Mass-specific rates of aerobic metabolism (/Mb) scale in inverse proportion to body mass (Mb). Thus, small hummingbirds display the highest / Mb known among vertebrates. Among all animals, higher /Mb values are known only in flying insects. The high body-mass-specific rates of metabolism seen in hummingbirds are made possible by high lung O2 diffusing capacities, cardiac outputs, ratios of capillary surface area to muscle fiber surface area, mitochondrial volume densities, cristae surface densities and concentrations of enzymes involved in energy metabolism. Current evidence from control analyses of O2 transport through the respiratory and cardiovascular systems and of metabolic fluxes through pathways of energy metabolism indicates shared control of maximum flux rates among multiple steps (i.e. the absence of single rate-limiting steps). This supports the suggestion that functional capacities at each step in linear pathways or processes are matched to each other, and provides an explanation for why the up-regulation of functional capacities has occurred at virtually all steps in the evolution of the smallest vertebrate homeotherms. Flying insects make use of a tracheal system for O2 transport and, like hummingbirds, possess a highly up-regulated biochemical machinery for substrate oxidation. Studies of hummingbirds and honeybees reveal closer matches between biochemical flux capacities and maximum physiological flux rates than in animals capable of lower maximum /Mb. It is proposed that the upper limits to functional capacities set the upper limit to /Mb. This upper limit to aerobic metabolic rate may contribute, along with other factors, towards establishing the lower limit to vertebrate homeotherm size.

Thoracic temperatures (Tth) of Libellula pulchella dragonflies during activity in the field were compared between age classes and with laboratory measures of optimal thoracic temperature for flight performance (Tth,opt; a trait that varies during adult maturation in this species). Newly emerged adults (tenerals) had mean Tth values during flight (34.5 °C; range 29–40 °C) that did not differ from their mean Tth,opt (34.6 °C; range 28.5–43.8 °C). Mature adults had higher and more precisely regulated thoracic temperatures (mean Tth 41.7 °C; range 37.5–45.2 °C), which were somewhat lower than their mean Tth,opt (43.6 °C; range 38.7–49.9 °C). Among matures, behaviors requiring the highest levels of flight exertion (aerial copulation; mate guarding; escalated territorial contests) caused an elevation of Tth above that of concurrently sampled individuals engaged in routine flight (mean Tth difference 1.3 °C), which raised mean Tth to a level that was not significantly different from Tth,opt (42.5 versus 43.5 °C). Compared with tenerals, matures spent more time flying, made longer-duration flights and showed a more restricted pattern of daily activity. Sympatric Anax junius dragonflies that regulate Tth endothermically had a uniform pattern of activity across the entire day, i.e. occupied a broader ecological niche than that of L. pulchella. These results support the hypotheses that optimal body temperature evolves to match the elevated body temperatures that occur during exercise and that the ecological benefits of an expanded niche are a secondary benefit rather than a primary selective force during the evolution of homeothermy and high body temperatures.

[Figure][1] Compared with endotherms, homeotherms have it easy: environmental temperature has little effect on our biological processes as we generate our own warmth internally. In contrast, ectotherms – which don't intrinsically maintain a constant body temperature – must constantly