Electrophysiological characteristics of cardiac function and the intensity of protein synthesis in cardiomyocytes during the arousal of susliks from hibernation

It has been demonstrated that during winter hibernation (body temperature 2-4 degrees C), the heart rate in ground squirrels is equal to 100 10-12 beats/min. At the initial stage of the arousal, while body temperature remains still low (9-10 degrees C), the heart rate may increase up to 160-200 beats/min. At this stage, practically all electrophysiological parameters of the heart correspond to those in active animals. These results may indicate the ability of "cold" heart in arousing ground squirrels to operate as a normothermic organ and reveal certain role of the heart in body warming. Significant increase of the intensity of protein synthesis in cardiomyocytes together with periodic changes in protein composition of their membranes were found during arousal which may account for regulation of the level of metabolism in cells and for adaptation of the latter to different temperatures.     

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.     

不同基础体温的家兔对热原检查结果的影响

对实验中使用的普通级家兔基础体温进行统计,观察不同基础体温家兔注射血液制品后的升温情况。按照中华人民共和国药典(2005)年版三部的热原检查规定进行测定,将实验家兔基础体温38.0℃~39.6℃分为3组:38.0℃~38.5℃为1组;38.6℃~39.0℃为2组;39.1℃~39.6℃为3组。注射制品后,家兔升温≥0.4℃记为升温家兔,统计升温≥0.4℃的家兔升温百分率并对其进行统计学处理。经卡方检验,升温家兔≥0.4℃的百分率1组与2组、1组与3组有显著性差异(p<0.05);2组与3组无显著性差异(p>0.05)。家兔对热原的敏感性随基础体温的高低而有明显的差异,基础体温偏低的家兔对热原更敏感,其升温幅度大于基础体温偏高的家兔。     

Effect of physical training on basal metabolism. (1). Aging and long-term exercise loaded at a moderate intensity in rats

Effect of training on basal metabolism in rats by means of long-term exercise loaded at a moderate intensity were studied. The Wistar-strain male rats were carefully bred at the room temperature of 23.0 +/- 1 degree C and humidity of 60%. The first physical training was carried out by motor driven treadmill for 8 weeks at a speed of 25 m/min for less than 15 min once daily and 6 times in a week after 4 weeks of birth. Continuously, the second training was carried out for 15 months at the same load of one time per week. Running ability and the recovery of glycogen in exhausted skeletal muscles on period of the first training, loaded from 4 weeks to 12 weeks after birth. There was no change on the basal metabolism between the trained and sedentary control rats. In general, the basal metabolism significantly fell by aging, for instance, 24 months rat's basal metabolism was 63% of 3-4 months rat's. The second training repressed a decline of running ability and rose the recovery of muscle glycogen in rats, though training could not be stopped lowering of basal metabolism in aged rats.     

THE INFLUENCE OF BROOD SIZE ON THE ENERGY METABOLISM AND WATER LOSS OF NESTLING GREAT TITS PARUS MAJOR MAJOR

At normal outdoor temperatures there is a distinct influence of brood size on the heat production of ten-day-old Great Tits. One ten-day-old nestling proved unable to maintain its body temperature at 12^°C. Two ten-day-old tits together in one nestbox at 12^Á°C were able to elevate the air temperature sufficiently to maintain homoiothermia. The same of course holds for tits in larger broods.
At an air temperature of 18^°C, six or seven ten-day-old tits placed in a nestbox elevated the air temperature to a level at which they almost reached a state of hyperthermia: their metabolism was at the basal level. The basal metabolism of a ten-day-old tit was found to be slightly more than 0–1800 kcal/h. The metabolism intensity of 12 tits in a nestbox at 12^°C was of the same order of magnitude.
Tits in broods comprising more than 12 or 13 nestlings at normal outdoor temperatures probably develop hyperthermia, which is unfavourable both for their energy and for their water balance.     

Reduction of metabolic rate and thermoregulation during daily torpor

Physiological mechanisms causing reduction of metabolic rate during torpor in heterothermic endotherms are controversial. The original view that metabolic rate is reduced below the basal metabolic rate because the lowered body temperature reduces tissue metabolism has been challenged by a recent hypothesis which claims that metabolic rate during torpor is actively downregulated and is a function of the differential between body temperature and ambient temperature, rather than body temperature per se. In the present study, both the steady-state metabolic rate and body temperature of torpid stripe-faced dunnarts, Sminthopsis macroura (Dasyuridae: Marsupialia), showed two clearly different phases in response to change of air temperature. At air temperatures between 14 and 30°C, metabolic rate and body temperature decreased with air temperature, and metabolic rate showed an exponential relationship with body temperature (r ²=0.74). The Q ₁₀ for metabolic rate was between 2 and 3 over the body temperature range of 16 to 32°C. The difference between body temperature and air temperature over this temperature range did not change significantly, and the metabolic rate was not related to the difference between body temperature and air temperature (P=0.35). However, the apparent conductance decreased with air temperature. At air temperatures below 14°C, metabolic rate increased linearly with the decrease of air temperature (r ²=0.58) and body temperature was maintained above 16°C, largely independent of air temperature. Over this air temperature range, metabolic rate was positively correlated with the difference between body temperature and air temperature (r ²=0.61). Nevertheless, the Q ₁₀ for metabolic rate between normothermic and torpid thermoregulating animals at the same air temperature was also in the range of 2–3. These results suggest that over the air temperature range in which body temperature of S. macroura was not metabolically defended, metabolic rate during daily torpor was largely a function of body temperature. At air temperatures below 14°C, at which the torpid animals showed an increase of metabolic rate to regulate body temperature, the negative relationship between metabolic rate and air temperature was a function of the differential between body temperature and air temperature as during normothermia. However, even in thermoregulating animals, the reduction of metabolic rate from normothermia to torpor at a given air temperature can also be explained by temperature effects.Abbreviations BM body mass - BMR basal metabolic rate - C apparent conductance - MR metabolic rate - RMR resting metabolic rate - RQ respiratory quotient - T _a air temperature - T _b body temperature - T _lc lower critical temperature - T _tc critical air temperature during torpor - TMR metabolic rate during torpor - TNZ thermoneutral zone - T difference between body temperature and air temperature - VO₂ rate of oxygen consumption     

Physiological convergence amongst ant-eating and termite-eating mammals

Ant- and termite-eating are among the few food habits common to monotremes, marsupials, and eutherians. Data are reported on the rate of metabolism and temperature regulation of 14 species of mammals having these food habits, including two monotremes, one marsupial and 11 eutherians. Small mammals with these habits have comparatively high body temperatures and high basal rates of metabolism, but ant- and termite-eaters that weigh more than 1 kg generally have low body temperatures and low basal rates of metabolism. The higher basal rates in small species ensure effective temperature regulation. Low body temperatures in large species principally result from low rates of metabolism. Rates of metabolism are low in these mammals because they use a food that has a limited availability and a low energy density, the density being further decreased in large species by the ingestion of non-nutritive material during feeding. Burrowing habits in some large species also contribute to low rates of metabolism. The combination of body size, food habits, and presence or absence of burrowing behaviour can account for all but about 6% of the range in basal rate in ant- and termite-eaters. Ants and termites, because of their locally clumped distributions, permit a larger mass in terrestrial predators than do other invertebrate prey. The reason why so many "primitive" mammals feed on ants and termites is that, once evolved, mammals with these habits are nearly impossible to displace ecologically, because much of ecological replacement is associated with high rates of reproduction, which are themselves correlated with high rates of metabolism in eutherians. Consequently, the ecological replacement of ant- and termite-eaters is inhibited, because this food habit does not permit high rates of metabolism, except at small masses.     

Effects of lifestyle, body composition, and physical fitness on cold tolerance in humans

In the present study, we attempted to clarify the effects of lifestyle and body compositions on basal metabolism and to clarify the effects of physical training on thermoregulatory responses to cold. Basal metabolism, body compositions, and questionnaires regarding lifestyle were evaluated in 37 students. From multiple linear regression analysis, sex, muscle weight, fat intake, and diurnal temperature were selected as significant explanatory variables. In a second experiment, rectal and the skin temperature at 7 different points as well as the oxygen uptake of eight males were measured at 10 degrees C for 90 min before and after training. The decline in rectal temperature that was observed before training was not observed after training. In addition, rectal temperature was significantly higher at post-training than at pre-training. These results suggest that some lifestyle factors affect cold tolerance; in particular, daily activity might improve our ability to control heat radiation and basal heat production.     

Effects of the menstrual cycle phase on the blood lactate responses to exercise

The effects of menstrual cycle phase on the blood lactate response to exercise were examined in eumenorrheic women (n=9). Exercise tests were performed at the mid-follicular and mid-luteal points in the menstrual cycle (confirmed by basal body temperature records and hormone levels). Blood lactates were measured at rest and during the recovery from exercise. Resting lactates were not different between the exercise tests; however, recovery lactates were significantly (p < 0.05) lower in the luteal compared to the follicular phase. The mechanism for these differences is unclear, but may be related to an estrogen mediated increased lipid metabolism inducing a concurrent reduction in carbohydrate metabolism. The present findings question the use of blood lactate monitoring as a suitable technique to measure exercise intensity in eumenorrheic women.     

Thermal energetics and torpor in the common pipistrelle bat, Pipistrellus pipistrellus (Vespertilionidae: Mammalia)

Rate of metabolism and body temperature were studied between -6°C and 38°C in the common pipistrelle bat Pipistrellus pipistrellus (Vespertilionidae), a European species lying close to the lower end of the mammalian size range (body mass 4.9±0.8g, N=28). Individuals maintained only occasionally a normothermic body temperature averaging 35.4±1.1°C (N=4) and often showed torpor during metabolic runs. The thermoneutral zone was found above 33°C, and basal rate of metabolism averaged 7.6±0.8mL O(2)h(-1) (N=28), which is 69% of the value predicted on the basis of body mass. Minimal wet thermal conductance was 161% of the expected value. During torpor, the rate of metabolism was related exponentially to body temperature with a Q(10) value of 2.57. Torpid bats showed intermittent ventilation, with the frequency of ventilatory cycles increasing exponentially with body temperature. Basal rate of metabolism (BMR) varied significantly with season and body temperature, but not with body mass. It was lower before the hibernation period than during the summer. The patterns observed are generally consistent with those exhibited by other vespertilionids of temperate regions. However, divergences occur with previous measurements on European pipistrelles, and the causes of the seasonal variation in BMR, which has only rarely been searched for among vespertilionids, remain to be examined.     

Rate of metabolism, temperature regulations, and evaporative water loss in the lesser gymnure Hylomys suillus (Insectivora, Mammalia)

Rate of metabolism, body temperature, wet thermal conductance, and evaporative water loss were measured at different ambient temperatures in four lesser gymnures Hylomys suillus. Gymnures responded as typical endothermic homoiotherms to changes in ambient temperature. Below the lower critical temperature of 32°C, they maintained a body temperature of 37.3± 0.3°C by an increased rate of metabolism. Minimum wet thermal conductance was 111% of that expected on the basis of body mass. Average basal rate of metabolism was 1.04 ml O₂ g⁻¹ h⁻¹, which represents 106% of the expected value. Within and above the thermoneutral zone, heat loss by evaporation did not account for more than 30% of the heat produced. As a consequence, the body temperature of gymnures was maintained 4°C above ambient temperature. These metabolic and thermoregulatory patterns differ strikingly from those of other members of the family Erinaceidac and can be interpreted as a result of physiological adaptation to a different ecology. Being smaller than hedgehogs and inhabiting montane tropical rainforests, lesser gymnures lack the physiological traits which enable many hedgehogs to invade hot, arid and/or strongly seasonal environments.     

Diving metabolism and thermoregulation in common and thick-billed murres

The diving and thermoregulatory metabolic rates of two species of diving seabrid, common (Uria aalge) and thick-billed murres (U. lomvia), were studied in the laboratory. Post-absorptive resting metabolic rates were similar in both species, averaging 7.8 W·kg^-1, and were not different in air or water (15–20°C). These values were 1.5–2 times higher than values predicted from published allometric equations. Feeding led to increases of 36 and 49%, diving caused increases of 82 and 140%, and preening led to increases of 107 and 196% above measured resting metabolic rates in common and thick-billed murres, respectively. Metabolic rates of both species increased linearly with decreasing water temperature; lower critical temperature was 15°C in common murres and 16°C in thick-billed murres. Conductance (assuming a constant body temperature) did not change with decreasing temperature, and was calculated at 3.59 W·m^-2·^oC^-1 and 4.68 W·m^-2·^oC^-1 in common and thick-billed murres, respectively. Murres spend a considerable amount of time in cold water which poses a significant thermal challenge to these relatively small seabirds. If thermal conductance does not change with decreasing water temperature, murres most likely rely upon increasing metabolism to maintain body temperature. The birds probably employ activities such as preening, diving, or food-induced thermogenesis to meet this challenge.Abbreviations ADL aerobic dive limit - BMR basal metabolic rate - FIT food-induced thermogenesis - MHP metabolic heat production - MR metabolic rate - PARR post-absorption resting rate - RMR resting metabolic rate - RQ respiratory quotient - SA surface area - STPD standard temperature and pressure (25°C, 1 ATM) - T _a ambient temperature - T _b body temperature - T _IC Iower critical temperatiure - TC thermal conductance - V oxygen consumption rate - W body mass     

Summer acclimatization in the short-tailed field vole,Microtus agrestis

We investigated the changes that occurred in basal and noradrenaline-induced metabolic rate, body temperature and body mass in short-tailed field voles,Microtus agrestis, during exposure to naturally increasing photoperiod and ambient temperature. These parameters were first measured in winter-acclimatized voles (n=8) and then in the same voles which had been allowed to seasonally acclimatize to photoperiod and ambient temperature (6 months later). Noradrenaline induced metabolic rate, basal metabolic rate and nonshivering thermogenesis were significantly higher in winter-acclimatized compared to summer-acclimatized voles. There was a significant positive relationship between basal metabolic rate and noradrenaline-induced metabolic rate. Body mass was significantly higher in summer-acclimatized compared to winter-acclimatized voles. There was a significant positive relationship between body mass and noradrenaline-induced metabolic rate in both winter-acclimalized and summer-acclimatized voles; however, there was no relationship between basal metabolic rate and body mass in either seasonal group of voles. Body temperature after measurements of basal metabolic rate was not significantly different in the seasonal cohorts of voles. However, body temperature was significantly higher in winter-acclimatized compared to summer-acclimatized voles after injection of noradrenaline. Previously we have found that a long photoperiod was not a sufficient stimulus to reduce thermogenic capacity in winter-acclimatized voles during cold exposure, since basal metabolic rate increased to compensate for a reduction in regulatory nonshivering thermogenesis. Here we found that a combination of increased ambient temperature and photoperiod did significantly reduce thermogenic capacity in winter-acclimatized voles. This provided evidence that the two aspects of non-shivering thermogenesis, obligatory and regulatory, are stimulated by different exogenous cues. Summer acclimatization in the shorttailed field vole is manifest as a significant decrease in both basal and noradrenaline-induced metabolic rate, combined with a significant increase in body mass.Abbreviations ANCOV A analysis of covariance - BAT brown adipose tissue - BM body mass - BMR basal metabolic rate - NST non-shivering thermogenesis - NA noradrenaline - V the maximum V recorded following mass specific injection of noradrenaline - V the maximum V recorded following mass specific injection of saline - T _a ambient temperature - T _b rectal body temperature - T _1c lower critical temperature - UCP uncoupling protein - V oxygen consumption     

Protein synthesis during acclimation of cold-blooded animals to various temperatures]

It was shown that the intensity of protein synthesis in cells of frogs, acclimated to 5 degrees C, is maintained at a high level, which is only 1.5-2 folds lower than that in animals acclimated to 20 C. In the process of acclimation to cold the intensity of synthesis decreases rapidly and already after 5 hours comprises one half of the value, which is characteristic of "warm" frogs, and the intensity of the process decreases more rapidly than the temperature of organs. On acclimation to warmth the intensity of protein synthesis increases and is getting stabilized at the level, characteristic of "warm" amphibia in 10-15 hours. It was shown that under various temperature conditions or conditions of acclimation specific proteins were synthesized against a background of the main groups of proteins.     

Food habits and the evolution of energetics in birds of paradise (Paradisaeidae)

Basal rates of metabolism, minimal thermal conductances, and body temperatures are reported for 13 species of birds of paradise that belong to nine genera. Body mass alone accounts for 91.7% of the variation in their basal rates. Basal rate in this family also correlates with food habits and the altitudinal limits to distribution. Species that feed almost exclusively on fruit have basal rates that average 79.4% of species in which >10% of the diet is insects, and species restricted to altitudes <1,000 m have basal rates that are 90.6% of those found at higher altitudes. The combination of body mass, food habits, and altitudinal distribution accounts for 99.0% of the variation in basal rate in the species studied. The application of food habits to a cladogram of the studied Paradisaeidae implies that frugivory and low basal rate were plesiomorphic in this family. The evolution of omnivory, defined as including >10% of the diet as insects, appears to have occurred at least twice, and in each case was associated with an increase in basal rate of metabolism. Basal rate increased at least thrice with a movement into the highlands. Basal rate, however, does not correlate with plumage dimorphism or with reproductive behavior. The basal rates of metabolism in manakins and birds of paradise, i.e., passerine frugivores, are greater than those found in nonpasserine frugivores. Thermal conductance correlates with body mass, which accounts for 85.8% of its variation in this family. Body temperature in paradisaeids, the mean of which was 40.2°C, may correlate with basal rate of metabolism.     

Energetics of East African Pycnonotids1

Rate of oxygen consumption was measured in five bulbuls (Family Pycnonotidae) from western Uganda to evaluate whether this group is indeed characterized by the very low basal rates of metabolism previously reported. For three of these species, body temperature and rate of metabolism were measured as a function of ambient temperature from 10°C to 35°C. In these species body temperature was highly variable, and declined with ambient temperature in Andropadus virens. Such variation, in conjunction with behavioral adjustments, may reduce heat loss at low ambient temperatures. Body mass accounted for 98 percent of the variation in the basal rates of metabolism presented here. Basal rates in these species ranged from 81 to 90 percent of values predicted by the Aschoff–Pohl relationship for passerines, whereas previous measurements ranged from 56 to 72 percent of predicted values. This difference may reflect differences in species or measurement techniques, which, if the latter, suggests that the reduction in metabolic rate in this family may be less than originally thought. These data underline the importance of continued data collection on the metabolism of tropical birds, few of which have been measured to date.     

Temperature-shock induction of multiple flagella induces additional synthesis of flagellum specific mRNAs and tubulin

Four mRNAs (alpha- and beta-tubulin, flagellar calmodulin and Class-I), specifically expressed when Naegleria amebae differentiate into flagellates, were followed at 5-10 min intervals during the temperature-shock induction of multiple flagella in order to better understand how basal body and flagellum number are regulated. Surprisingly, tubulin synthesis continued during the 37 min temperature shock. An initial rapid decline in alpha- and beta-tubulin and flagellar calmodulin mRNAs was followed by a rapid re-accumulation of mRNAs before the temperature was lowered. mRNA levels continued to increase until they exceeded control levels by 4-21%. Temperature shock delayed flagella formation 37 min, produced twice as much tubulin protein synthesis and three fold more flagella. Labeling with an antibody against Naegleria centrin suggested that basal body formation was also delayed 30-40 min. An extended temperature shock demonstrated that lowering the temperature was not required for return of mRNAs to near control levels suggesting that induction of multiple flagella and the formation of flagella per se are affected in different ways. We suggest that temperature-shock induction of multiple flagella reflects increased mRNA accumulation combined with interference with the regulation of the recently reported microtubule-nucleating complex needed for basal body formation.     

Warm-up rates during arousal from torpor in heterothermic mammals: physiological correlates and a comparison with heterothermic insects

Summary This study examines the relationship between warm-up rate, body mass, metabolic rate, thermal conductance and normothermic body temperature in heterothermic mammals during arousal from torpor. Predictions based on the assumption that the energetic cost of arousal has been minimised are tested using data for 35 species. The observation that across-species warm-up rate correlates negatively with body mass is confirmed using a comparative technique which removes confounding effects due to the non-independence of species data due to shared common ancestry. Mean warm-up rate during arousal correlates negatively with basal metabolic rate and positively with the temperature difference through which the animal warms, having controlled for other factors. These results suggest that selection has operated to minimise the overall energetic, cost of warm-up. In contrast, peak warm-up rate during arousal correlates positively with peak metabolic rate during arousal, and negatively with thermal conductance, when body mass has been taken into account. These results suggest that peak warm-up rate is more sensitive to the fundamental processes of heat generation and loss. Although heterothermic marsupials have lower normothermic body temperatures and basal metabolic rates, marsupials and heterothermic eutherian mammals do not differ systematically in warm-up rate. Pre-flight warm-up rates in one group of endothermic insects, the bees, are significantly higher than predictions based on rates of arousal of a mammal of the same body mass.Abbreviations BMR basal metabolic rate - ICM independent comparisons method - MWR mean warm-up rate - PMR peak metabolic rate - PWR peak·warm-up rate - Tb_activity body temperature during activity - Tb_torpor body temperature during torpor - T _arousal increase in body temperature during arousal     

The energetics of New Zealand's ducks

Measurements on rates of metabolism and temperature regulation are presented from nine populations of seven species of ducks resident in New Zealand. An analysis of these data and those from 18 additional species obtained from the literature indicates that basal rate of metabolism in anatids correlates with body mass and restriction to the Australian-New Zealand region: these 'southern' species have basal rates that average 70% of those from the Northern Hemisphere. The low basal rates of southern anatids may reflect reduced pectoral muscle masses in association with the absence of migratory habits and/or life on land masses without eutherian predators. New Zealand flightless teal (Anas aucklandica nesiotis, Anas aucklandica aucklandica) do not have mass-independent basal rates that differ from those found in flighted ducks living in the same region, although flightless teal have lower total basal rates than most ducks as a result of small masses. Minimal thermal conductance in this sample is determined by body mass alone. Regulated body temperature is negatively correlated with body mass.     

The thermal and energetic significance of clustering in the speckled mousebird,Colius striatus

Summary The effect of clustering behaviour on metabolism, body temperature, thermal conductance and evaporative water loss was investigated in speckled mousebirds at temperatures between 5 and 36°C. Within the thermal neutral zone (approximately 30–35 °C) basal metabolic rate of clusters of two birds (32.5 J·g^-1·h^-1) and four birds (28.5 J·g^-1·h^-1) was significantly lower by about 11% and 22%, respectively, than that of individuals (36.4 J·g^-1·h^-1). Similarly, below the lower critical temperature, the metabolism of clusters of two and four birds was about 14% and 31% lower, respectively, than for individual birds as a result of significantly lower total thermal conductance in clustered birds. Body temperature ranged from about 36 to 41°C and was positively correlated with ambient temperature in both individuals and clusters, but was less variable in clusters. Total evaporative water loss was similar in individuals and clusters and averaged 5–6% of body weight per day below 30°C in individuals and below 25°C in clusters. Above these temperatures total evaporative water loss increased and mousebirds could dissipate between 80 and 90% of their metabolic heat production at ambient temperatures between 36 and 39°C. Mousebirds not only clustered to sleep between sunset and sunrise but were also observed to cluster during the day, even at high ambient temperature. Whereas clustering at night and during cold, wet weather serves a thermoregulatory function, in that it allows the brrds to maintain body temperature at a reduced metabolic cost, clustering during the day is probably related to maintenance of social bonds within the flock.Abbreviations BMR basal metabolic rate - bw body weight - C _totab total thermal conductance - EWI evaporative water loss - M metabolism - RH relative humidity - T _a ambient temperature - T _b body temperature - T _ch chamber temperature - T _cl cluster temperature - TEWL total evaporative water loss - LCT lower critical temperature - TNZ thermal neutral zone