Seasonal changes in the preferred body temperature, cardiovascular, and respiratory responses to hypoxia in the toad, Bufo paracnemis

Estivation is accompanied by a reduction of oxygen consumption in amphibians during drought. We tested the hypothesis that, during the dry season, the toad Bufo paracnemis selects a lower preferred body temperature (T(b)), and would be less sensitive to hypoxia, than during its active period. Therefore, during winter (dry season in S?o Paulo state, Brazil) and summer, we measured the effects of hypoxia (7% inspired O(2)) on preferred T(b). Additionally, pulmonary ventilation, heart rate, blood pressure, and oxygen consumption were also measured in toads at 15 and 25 degrees C. Blood gases were measured at 25 degrees C. Oxygen consumption was significantly higher during summer in toads at 25 degrees C. Under normoxia, preferred T(b) was higher during summer than during winter, and hypoxia caused a drop in preferred T(b) during both seasons. In both seasons, toads at 15 degrees C showed reduced pulmonary ventilation, heart rate, and blood pressure, and hypoxia had no effect. At 25 degrees C during summer only, hypoxia caused an increase in ventilation. Season had no effect on blood gases. We conclude that B. paracnemis displays an endogenous seasonal pattern of thermoregulation and control of ventilation. The decreased preferred T(b) and the physiological responses to hypoxia may be beneficial to toads encountering drought and when food is not available.     

Ingested energy differs between populations of the toad Bufo bankorensis from different altitudes

We measured ingested energy (E(i)) and apparent digestibility efficiency (ADE) in two populations of Bufo bankorensis from different altitudes at three temperatures and during two seasons to test the hypothesis that the optimal temperature range (T(opt)) for E(i) and ADE has shifted to the lower range in highland toads and winter toads. The T(opt) for E(i) was 22 degrees C for the lowland and highland toads and did not vary between seasons, thus falsifying the hypothesis. ADE of the toads was 96%-99% at 15 degrees -30 degrees C, and there was no difference between populations or seasons. Furthermore, when fed with fast-moving prey, the toads from both altitudes had similarly low E(i) at 15 degrees C; when fed with slow-moving prey, the highland toads increased E(i) at 15 degrees C, but the lowland toads did not. These results suggest that the toads from different altitudes had different appetites, even though their feeding locomotion was hampered in both populations at low temperatures.     

Seasonal changes in the cardiorespiratory responses to hypercarbia and temperature in the bullfrog, Rana catesbeiana

We assessed the seasonal variations in the effects of hypercarbia (3 or 5% inspired CO2) on cardiorespiratory responses in the bullfrog Rana catesbeiana at different temperatures (10, 20 and 30 degrees C). We measured breathing frequency, blood gases, acid-base status, hematocrit, heart rate, blood pressure and oxygen consumption. At 20 and 30 degrees C, the rate of oxygen consumption had a tendency to be lowest during winter and highest during summer. Hypercarbia-induced changes in breathing frequency were proportional to body temperature during summer and spring, but not during winter (20 and 30 degrees C). Moreover, during winter, the effects of CO2 on breathing frequency at 30 degrees C were smaller than during summer and spring. These facts indicate a decreased ventilatory sensitivity during winter. PaO2 and pHa showed no significant change during the year, but PaCO2 was almost twice as high during winter than in summer and spring, indicating increased plasma bicarbonate levels. The hematocrit values showed no significant changes induced by temperature, hypercarbia or season, indicating that the oxygen carrying capacity of blood is kept constant throughout the year. Decreased body temperature was accompanied by a reduction in heart rate during all four seasons, and a reduction in blood pressure during summer and spring. Blood pressure was higher during winter than during any other seasons whereas no seasonal change was observed in heart rate. This may indicate that peripheral resistance and/or stroke volume may be elevated during this season. Taken together, these results suggest that the decreased ventilatory sensitivity to hypercarbia during winter occurs while cardiovascular parameters are kept constant.     

Temperature-induced changes in the locomotor capacity of juveniles of Marenzelleria viridis (Polychaeta, Spionidae)

Abstract. Populations of Marenzelleria viridis in the Chester River (Kent County, Maryland) experience temperatures ranging from over 30°C in summer to near freezing in winter. Interestingly, M. viridis swims actively in winter. This observation led us to examine the relationship between locomotor capacity and temperature in individuals of M. viridis . Juvenile specimens were collected in February ("cold animals") and June ("warm animals"). Video analysis revealed that swimming is achieved by flexing the body in cyclic, helical waves. Wave frequencies were measured as an index of locomotor capacity at 5°C, 15°C, and 25°C. The mean wave frequencies of cold animals were 5.4 Hz at 5°C and 7.1 Hz at 15°C (Q₁₀= 1.3); the mean wave frequencies of warm animals were 6.1 Hz at 15°C and 7.8 Hz at 25°C (Q₁₀= 1.3). The effects of changes in water viscosity on wave frequency between 5–25°C were not significant. These results demonstrate that the temperature sensitivity of locomotor capacity in juvenile M. viridis is quite low. We conclude that low temperature sensitivity enables M. viridis to be active throughout the year.     

Thermotropic behavior of major phospholipids from marine invertebrates: changes with warm-acclimation and seasonal acclimatization

The crystal-liquid crystal-isotropic melt phase transitions of phosphatidylcholine (PC) and phosphatidylethanolamine (PE) from muscle tissue of five species (actinia Metridium senile fimbriatum, mussel Crenomytilus grayanus, sea-urchin Strongylocentrotus intermedius, starfish Distolasterias nipon and the ascidian Halocynthia aurantium) of marine invertebrates, collected in winter at 0 degrees C and then acclimated to 18.5 degrees C for 5 days, were studied by differential scanning calorimetry and polarising microscopy. To elevate temperature from 0 to 18.5 degrees C, we used the rate of 4.5 degrees C/h. Although phase transitions of both phospholipids from animals collected in summer occurred already at temperatures below -1.7 oC (minimal temperature of seawater in winter), compensatory mechanisms resulted in a decrease by 29-43 oC in the phase transition temperature of PE in winter. Thermotropic behavior of PCs changed in various trends. However, the total heat of their phase transitions always decreased in winter compared with summer. For all species, except the mussel, the time of warm-acclimation was insufficient to adjust the thermotropic behavior of either phospholipid. Nevertheless, the unsaturation index decreased to achieve summer values, due primarily to decreased proportions of eicosapentaenate and docosahexaenate. The accumulation of arachidonate, during warm-acclimation, might be connected to the signalling properties of n-6 eicosanoids. Absence of effective homeoviscous mechanisms suggests that most of the studied marine invertebrates have very limited capacity to survive an acute temperature elevation, e.g. at the appearance of thermal currents.     

Oxygen consumption of a terrestrial toad (Bufo viridis) and semi-aquatic frog (Rana ridibunda)

1. Oxygen consumption was measured before and after dehydration at different ambient temperatures (Ta) in the terrestrial toad Bufo viridis and the semi-aquatic frog Rana ridibunda. 2. The metabolic rates at Tas between 14 and 27 degrees C were almost the same for R. ridibunda and B. viridis. 3. The metabolic rate at higher Tas (between 27 and 36 degrees C) was higher in R. ridibunda than in B. viridis. This situation was found before and after dehydration. 4. A similar situation was found with CO2 production, which was higher at high Tas in R. ridibunda compared with B. viridis.     

Effects of season, temperature, and body mass on the standard metabolic rate of tegu lizards (Tupinambis merianae)

Abstract This study examined how the standard metabolic rate of tegu lizards, a species that undergoes large ontogenetic changes in body weight with associated changes in life-history traits, is affected by changes in body mass, body temperature, season, and life-history traits. We measured rates of oxygen consumption (Vo(2)) in 90 individuals ranging in body mass from 10.4 g to 3.75 kg at three experimental temperatures (17 degrees , 25 degrees , and 30 degrees C) over the four seasons. We found that standard metabolic rate scaled to the power of 0.84 of body mass at all experimental temperatures in all seasons and that thermal sensitivity of metabolism was relatively low (Q(10) approximately 2.0-2.5) over the range from 17 degrees to 30 degrees C regardless of body size or season. Metabolic rates did vary seasonally, being higher in spring and summer than in autumn and winter at the same temperatures, and this was true regardless of animal size. Finally, in this study, the changes in life-history traits that occurred ontogenetically were not accompanied by significant changes in metabolic rate.     

Density regulation in toad populations (Epidalea calamita,Bufotes viridis) by differential winter survival of juveniles

The size of amphibian populations varies considerably between years, so that systematic trends in dynamics are difficult to detect. Informed conservation management of presumably declining populations requires the identification of the most sensitive life stage. In temperate-zone anurans there is growing evidence that juveniles hibernating for the first time suffer from substantial winter losses. In two syntopic toads (Epidalea calamita, Bufotes viridis) we monitored survival of such juveniles during four consecutive winters in the natural habitat and in four temperature treatments (3°, 5 °C, 10°/15 °C or 20 °C, natural light-dark cycle) in temperature-controlled chambers during winter. Specifically, we tested the hypotheses that (1) winter mortality of juvenile toads which hibernate for the first time in their life is an important component of population dynamics, and that (2) mortality rates differed between the two species. Parameters quantified were size-dependent winter mortality and body condition of pre- and post-hibernating juveniles. Field data provided evidence for the important role of winter mortality of first-hibernators in population dynamics. Choice of hibernacula differed in E. calamita between small and medium-sized individuals and also between the two species suggesting distinct mortality risks. The inability of small E. calamita to reach frost-proof hibernacula by burrowing, and the exposure of small B. viridis to predators are the most probable causes of size-assortative winter mortality. In conclusion, E. calamita juveniles may benefit from rising average winter temperatures in the future by decreased risk of freezing to death, whereas predator-caused winter mortality of B. viridis juveniles will also depend on the effects of climate warming on predator phenology.     

A comparative analysis of the temperature response of leaf elongation in Bromus stamineus and Lolium perenne plants in the field: intrinsic and size-mediated effects

BACKGROUND AND AIMS: Growth of grass species in temperate-humid regions is restricted by low temperatures. This study analyses the origin (intrinsic or size-mediated) and mechanisms (activity of individual meristems vs. number of active meristems) of differences between Bromus stamineus and Lolium perenne in the response of leaf elongation to moderately low temperatures. METHODS: Field experiments were conducted at Balcarce, Argentina over 2 years (2003 and 2004) using four cultivars, two of B. stamineus and two of L. perenne. Leaf elongation rate (LER) per tiller and of each growing leaf, number of growing leaves and total leaf length per tiller were measured on 15-20 tillers per cultivar, for 12 (2003) or 10 weeks (2004) during autumn and winter. KEY RESULTS: LER was faster in B. stamineus than in L. perenne. In part, this was related to size-mediated effects, as total leaf length per tiller correlated with LER and B. stamineus tillers were 71% larger than L. perenne tillers. However, accounting for size effects revealed intrinsic differences between species in their temperature response. These were based on the number of leaf meristems simultaneously active and not on the (maximum) rate at which individual leaves elongated. Species differences were greater at higher temperatures, being barely notable below 5 degrees C (air temperature). CONCLUSIONS: Bromus stamineus can sustain a higher LER per tiller than L. perenne at air temperatures > 6 degrees C. In the field, this effect would be compounded with time as higher elongation rates lead to greater tiller sizes.     

Seasonal patterns in body temperature of free-living rock hyrax (Procavia capensis)

Rock hyrax (Procavia capensis) are faced with large daily fluctuations in ambient temperature during summer and winter. In this study, peritoneal body temperature of free-living rock hyrax was investigated. During winter, when low ambient temperatures and food supply prevail, rock hyrax maintained a lower core body temperature relative to summer. In winter body temperatures during the day were more variable than at night. This daytime variability is likely a result of body temperatures being raised from basking in the sun. Body temperatures recorded during winter never fell to low levels recorded in previous laboratory studies. During summer ambient temperatures exceeded the thermoneutral zone of the rock hyrax throughout most of the day, while crevice temperatures remained within the thermoneutral zone of rock hyrax. However, in summer variation in core body temperature was small. Minimum and maximum body temperatures did not coincide with minimum and maximum ambient temperatures. Constant body temperatures were also recorded when ambient temperatures reached lethal limits. During summer it is likely that rock hyrax select cooler refugia to escape lethal temperatures and to prevent excessive water loss. Body temperature of rock hyrax recorded in this study reflects the adaptability of this animal to the wide range of ambient temperatures experienced in its natural environment.     

Estimation of base temperatures for nine weed species

Experiments were conducted to test several methods for estimating low temperature thresholds for seed germination. Temperature responses of nine weeds common in annual agroecosystems were assessed in temperature gradient experiments. Species included summer annuals (Amaranthus albus, A. palmeri, Digitaria sanguinalis, Echinochloa crus-galli, Portulaca oleracea, and Setaria glauca), winter annuals (Hirschfeldia incana and Sonchus oleraceus), and Conyza canadensis, which is classified as a summer or winter annual. The temperature below which development ceases (Tbase) was estimated as the x-intercept of four conventional germination rate indices regressed on temperature, by repeated probit analysis, and by a mathematical approach. An overall Tbase estimate for each species was the average across indices weighted by the reciprocal of the variance associated with the estimate. Germination rates increased linearly with temperature between 15 degrees C and 30 degrees C for all species. Consistent estimates of Tbase were obtained for most species using several indices. The most statistically robust and biologically relevant method was the reciprocal time to median germination, which can also be used to estimate other biologically meaningful parameters. The mean Tbase for summer annuals (13.8 degrees C) was higher than that for winter annuals (8.3 degrees C). The two germination response characteristics, Tbase and slope (rate), influence a species' germination behaviour in the field since the germination inhibiting effects of a high Tbase may be offset by the germination promoting effects of a rapid germination response to temperature. Estimates of Tbase may be incorporated into predictive thermal time models to assist weed control practitioners in making management decisions.     

The thermal physiology of two sympatric treefrogs Hyla cinerea and Hyla chrysoscelis (Anura; Hylidae)

Metabolic rates, temperature acclimation, lipid deposition and temperature tolerance were investigated in two species of hylid treefrogs, the green treefrog (Hyla cinerea) and the coastal plain (Cope's) gray treefrog (Hyla chrysoscelis). The rate of oxygen consumption at rest differed between the two species only at 30 degrees C; there was no difference in respiratory metabolism at lower ambient temperatures. Hyla cinerea generally completed metabolic acclimation earlier than H. chrysoscelis, particularly at high temperatures; both species appeared to be fully acclimated in 6 days or less. The gray treefrog is less tolerant of high ambient temperatures than the green treefrog; mean upper lethal temperature was 41.5 degrees C for Hyla chrysoscelis and 43.7 degrees C for H. cinerea. Metabolized energy was higher at high ambient temperatures (i.e. 29 degrees C) for H. chrysoscelis than H. cinerea, while the reverse was true at 19 degrees C. The coefficient of utilization (100 X metabolized energy/gross energy intake) did not vary significantly between species or within species over the ambient temperature range of 19-24 degrees C; H. chrysoscelis had a significantly higher efficiency at 29 degrees C. Lipid reserves were generally similar in the two species throughout the summer. Differences in behavior, seasonal variation in activity and timing of reproduction are all related to thermal physiology and may play a role in determining the distributional limits of the two species.     

Low temperature cardiac response to exhaustive exercise in fish with different levels of winter quiescence

We examined the cardiac responses of different fish species to anaerobic exercise at low temperatures (3 degrees C). Three species of sympatric warmwater fish with perceived differences in winter activity were used for this comparative study: the winter-quiescent largemouth bass (Micropterus salmoides); the winter-active white bass (Morone chrysops); and the intermediately winter-active black crappie (Pomoxis nigromaculatus). Perceived differences in winter activity were reflected in cardiac responses; e.g. basal cardiac values were lowest for largemouth bass, highest for white bass, and intermediate for black crappie. In addition, cardiac recovery was most rapid for white bass, slowest for largemouth bass and intermediate for black crappie. When disturbed at low temperatures, largemouth bass and black crappie elevated cardiac output principally through increases in heart rate despite substantial decreases in stroke volume. Conversely, white bass principally used stroke volume modulation to change cardiac output. The results of this study indicate that different species respond differently to exercise at low temperatures. Management strategies should recognize that such variation exists and ensure that management decisions are based upon an understanding of the low temperature exercise physiology and winter biology of the species of interest.     

Variation of microsatellites BM224 and Bcal7 in populations of green toads (Bufo viridis complex) with various nuclear DNA content and ploidy

We studied variation of microsatellites BM224 and Bcal7 in three species of the Bufo viridis diploid-polyploid complex. We found that locus Bcal7 in all examined samples was monomorphic. Three alleles of microsatellite BM224 were found. Among tetraploid toads, the western species B. oblongus had only one allele variant, whereas the eastern species B. pewzowi had two other alleles. Similar distribution of alleles was observed in triploid specimens, collected in the area borders of tetraploid and diploid species. Among samples of diploid toad B. viridis, we found all three allele variants of microsatellite BM224. Their distribution was geographically determined. A comparison of allele distribution with genome size variation in diploid toads showed very similar patterns.     

Temperature dependent characteristics of intestinal glycyl-L-leucine dipeptidase in boreal zone fish

Three kinds of boreal zone fish were investigated for gastrointestinal glycyl-L-leucine (GL) dipeptide cleaving activity as a function of feeding stage and seasonal changes. The enzyme activity tested in the perch (Perca fluviatilis L.) intestine increased steadily during digestion and rapidly disappeared after completion. The temperature characteristics and the seasonal changes in dipeptide cleaving activity in pike perch (Stizostedion lucioperca L.) and bream (Abramis brama L.) were studied. In summer, the maximal activities in the pike perch and the bream were found at temperatures of 40 and 30 degrees C, respectively. In winter, the temperature of maximal activity in pike perch fell to only 30 degrees C, whereas no changes were observed in bream. The activation energies in bream and pike perch were several times lower in winter than in summer. Seasonal changes in the dipeptide cleaving activity at low temperature relative to that at the temperature of maximal activity were found. At high temperatures, the stability of the enzyme decreases in winter and increases in summer, but in the presence of a substrate the thermal stability of the enzyme increases both in winter and in summer. In our experiments, we found that in these fish, GL dipeptidase was unstable at 0 and -10 degrees C.     

Seasonal changes in thermal responses of urban residents to cold exposure

To determine whether urban circumpolar residents show seasonal acclimatisation to cold, thermoregulatory responses and thermal perception during cold exposure were examined in young men during January-March (n=7) and August-September (n=8). Subjects were exposed for 24 h to 22 and to 10 degrees C. Rectal (T(rect)) and skin temperatures were measured throughout the exposure. Oxygen consumption (VO(2)), finger skin blood flow (Q(f)), shivering and cold (CDT) and warm detection thresholds (WDT) were assessed four times during the exposure. Ratings of thermal sensations, comfort and tolerance were recorded using subjective judgement scales at 1-h intervals. During winter, subjects had a significantly higher mean skin temperature at both 22 and 10 degrees C compared with summer. However, skin temperatures decreased more at 10 degrees C in winter and remained higher only in the trunk. Finger skin temperature was higher at 22 degrees C, but lower at 10 degrees C in the winter suggesting an enhanced cold-induced vasoconstriction. Similarly, Q(f) decreased more in winter. The cold detection threshold of the hand was shifted to a lower level in the cold, and more substantially in the winter, which was related to lower skin temperatures in winter. Thermal sensations showed only slight seasonal variation. The observed seasonal differences in thermal responses suggest increased preservation of heat especially in the peripheral areas in winter. Blunted vasomotor and skin temperature responses, which are typical for habituation to cold, were not observed in winter. Instead, the responses in winter resemble aggravated reactions of non-cold acclimatised subjects.     

Developmental plasticity of mating calls enables acoustic communication in diverse environments

Male calls of the katydid Neoconocephalus triops exhibit substantial developmental plasticity in two parameters: (i) calls of winter males are continuous and lack the verse structure of summer calls and (ii) at equal temperatures, summer males produce calls with a substantially higher pulse rate than winter males. We raised female N. triops under conditions that reliably induced either summer or winter phenotype and tested their preferences for the call parameters that differ between summer and winter males.Neither generation was selective for the presence of verses, but females had strong preferences for pulse rates: only a narrow range of pulse rates was attractive. The attractive ranges did not differ between summer and winter females. Both male pulse rate and female preference for pulse rate changed with ambient temperature, but female preference changed more than the male calls.As a result, the summer call was attractive only at 25 degrees C, whereas the slower winter call was attractive only at 20 degrees C. Thus, developmental plasticity of male calls compensates for differences in temperature dependency between calls and preferences and enables the communication system to function in heterogeneous environments. The potential role of call plasticity during the invasion of new habitats is discussed.     

Mitochondrial function in seasonal acclimatization versus latitudinal adaptation to cold in the lugworm Arenicola marina (L.)

Previous studies in marine ectotherms from a latitudinal cline have led to the hypothesis that eurythermal adaptation to low mean annual temperatures is energetically costly. To obtain more information on the trade-offs and with that the constraints of thermal adaptation, mitochondrial functions were studied in subpolar lugworms (Arenicola marina L.) adapted to summer cold at the White Sea and were compared with those in boreal specimens from the North Sea, either acclimatized to summer temperatures or to winter cold. During summer, a comparison of mitochondria from subpolar and boreal worms revealed higher succinate oxidation rates and reduced Arrhenius activation energies (Ea) in state 3 respiration at low temperatures, as well as higher proton leakage rates in subpolar lugworms. These differences reflect a higher aerobic capacity in subpolar worms, which is required to maintain motor activity at low but variable environmental temperatures--however, at the expense of an elevated metabolic rate. The lower activity of citrate synthase (CS) found in subpolar worms may indicate a shift in metabolic control within mitochondria. In contrast, acclimatization of boreal lugworms to winter conditions elicited elevated mitochondrial CS activities in parallel with enhanced mitochondrial respiration rates. With falling acclimation temperatures, the significant Arrhenius break temperature in state 3 respiration (11 degrees C) became insignificant (5 degrees C) or even disappeared (0 degrees C) at lower levels of Arrhenius activation energies in the cold, similar to a phenomenon known from hibernating vertebrates. The efficiency of aerobic energy production in winter mitochondria rose as proton leakage in relation to state 3 decreased with cold acclimation, indicated by higher respiratory control ratio values and increased adenosine diphosphate/oxygen (ADP/O) ratios. These transitions indicate reduced metabolic flexibility, possibly paralleled by a loss in aerobic scope and metabolic depression during winter cold. Accordingly, these patterns contrast those found in summer-active, cold-adapted eurytherms at high latitudes.     

Temperature and adrenoceptors in the frog heart

1. Cardiac adrenergic receptors in a frog, Rana tigrina, were examined in winter and summer months using isolated atria preparation maintained at 24 degrees, 14 degrees and 6 degrees C. Treatments included an examination of the atrial responses to selective alpha and beta adrenergic agonists (phenylephrine and isoproterenol respectively) and antagonists (phentolamine and propranolol). 2. Basal atrial beating rates differed between summer and winter months and increased with temperature. 3. Phenylephrine produced dose-dependent increases in the atrial beating rate and tension in the winter frogs only at 6 degrees C. These increases were blunted by phentolamine. 4. Isoproterenol produced positive chronotropic effects of 14 degrees and 24 degrees C but not at 6 degrees C in both summer and winter frogs; these effects were abolished by propranolol. Further, at 6 degrees C, the contractile response of the atrial tissue to isoproterenol was very sensitive. 5. Data suggests that the alpha adrenoceptor might be physiologically important to the frog in the low temperature environment of the cold season, during which period the cardiac beta adrenergic activity would be minimal or even absent.     

Seasonal changes in daily metabolic patterns of tegu lizards (Tupinambis merianae) placed in the cold (17 degrees C) and dark

Abstract Oxygen consumption rate was measured continuously in young tegu lizards Tupinambis merianae exposed to 4 d at 25 degrees C followed by 7-10 d at 17 degrees C in constant dark at five different times of the year. Under these conditions, circadian rhythms in the rate of oxygen consumption persisted for anywhere from 1 d to the entire 2 wk in different individuals in all seasons except the winter. We also saw a progressive decline in standard oxygen consumption rate (at highly variable rates in different individuals) to a very low rate that was seasonally independent (ranging from 19.1 +/- 6.2 to 27.7 +/- 0.2 mL kg(-1) h(-1) across seasons). Although this degree of reduction appeared to take longer to invoke when starting from higher metabolic rates, tegu lizards reduced their metabolism to the low rates seen in winter dormancy at all times of the year when given sufficient time in the cold and dark. In the spring and summer, tegus reduced their standard metabolic rate (SMR) by 80%-90% over the experimental run, but only roughly 20%-30% of the total fall was due to the reduction in temperature; 70%-80% of the total fall occurred at constant temperature. By autumn, when the starting SMR on the first night at 25 degrees C was already reduced by 59%-81% (early and late autumn, respectively) from peak summer values, virtually all of the fall (63%-83%) in metabolism was due to the reduction in temperature. This suggests that the temperature-independent reduction of metabolism was already in place by autumn before the tegus had entered winter dormancy.