Can birds do it too? Evidence for convergence in evaporative water loss regulation for birds and mammals.

Birds have many physiological characteristics that are convergent with mammals. In the light of recent evidence that mammals can maintain a constant insensible evaporative water loss (EWL) over a range of perturbing environmental conditions, we hypothesized that birds might also regulate insensible EWL, reflecting this convergence. We found that budgerigars (Melopsittacus undulatus) maintain EWL constant over a range of relative humidities at three ambient temperatures. EWL, expressed as a function of water vapour pressure deficit, differed from a physical model where the water vapour pressure deficit between the animal and the ambient air is the driver of evaporation, indicating physiological control of EWL. Regulating EWL avoids thermoregulatory impacts of varied evaporative heat loss; changes in relative humidity had no effect on body temperature, metabolic rate or thermal conductance. Our findings that a small bird can regulate EWL are evidence that this is a common feature of convergently endothermic birds and mammals, and may therefore be a fundamental characteristic of endothermy.

Environmental physiology of a small marsupial inhabiting arid floodplains.

Giles' planigale (Planigale gilesi) is among the smallest extant marsupials and inhabits deep soil cracks in arid floodplains. We examined whether its physiology shows specific adaptations to its extreme habitat. Metabolic rate, body temperature, evaporative water loss and thermal conductance were measured for eight planigales (average mass 9 g) exposed to four different ambient temperatures ranging from 10 degrees C to 32 degrees C. Water economy and respiratory variables were measured for the first time in this species. All of these standard physiological variables conformed to allometrically-predicted values for a marsupial. All variables were significantly affected by ambient temperature, except tidal volume and dry thermal conductance. Metabolic rate increased substantially at low ambient temperatures, as required to maintain a relatively constant body temperature of about 32-34 degrees C. This increased oxygen demand was accommodated by increased ventilation rather than increased oxygen extraction. Planigales had a comparatively high point of relative water economy of 19.1 degrees C, consistent with their small body size and arid habitat. Torpor reduced energy expenditure by 79% and evaporative water loss by 62%. Our study suggests that torpor use, along with behavioural adaptations, suffice for P. gilesi to live underground in arid habitats without further physiological adaptations.

Metabolic, ventilatory and hygric physiology of the chuditch (Dasyurus geoffroii; Marsupialia, Dasyuridae).

The chuditch is a large carnivorous dasyurid marsupial. Historically it had one of the widest geographical distributions of all marsupials, encompassing much of arid Australia, but it is now restricted to the mesic south-west of Western Australia. It is therefore of interest to determine if its physiology better reflects adaptation to its historically arid or present mesic habitat. The basic physiological parameters of the chuditch conform to other marsupials. Body mass of males (1385 g) was >400% of that predicted by phylogeny and this may be related to its carnivorous diet. Body temperature was 33.9 degrees C at ambient temperatures < or = thermoneutrality, with hyperthermia occurring above thermoneutrality. Basal metabolic rate was 0.361 mL O(2) g(-1) h(-1) at an ambient temperature of 31 degrees C. Metabolic rate increased below the thermoneutral zone by 0.038 mL O(2) g(-1) h(-1) degrees C(-1), and above the thermoneutral zone to 0.444+/-0.059 mL O(2) g(-1) h(-1) at 33.3 degrees C. Standard evaporative water loss was 0.498+/-0.071 mg g(-1) h(-1) at an ambient temperature of 26.0 degrees C, and increased at higher ambient temperatures due to panting and licking. Changes in wet thermal conductance largely reflected changes in evaporative heat loss, and dry thermal conductance increased at high ambient temperature due in part to posture change. Ventilatory parameters were consistent with metabolic demands in and below thermoneutrality, and suggested augmented evaporative heat loss above the thermoneutral zone. Chuditch had a high point of relative water economy of 22.6 degrees C, indicating favourable water economy at even moderate ambient temperatures, due to its low evaporative water loss rather than high metabolic water production. Chuditch were physiologically more similar to marsupials from arid rather than mesic habitats, better reflecting their historical distribution than their current geographical range.

Environmental correlates of physiological variables in marsupials.

We analyzed body temperature (T(b)), basal metabolic rate (BMR), wet thermal conductance (C(wet)), and evaporative water loss (EWL) of marsupials by conventional and phylogenetically corrected regression. Allometric effects were substantial for BMR, C(wet), and EWL but not T(b). There was a strong phylogenetic signal for mass and all physiological traits. A significant phylogenetic signal remained for BMR, C(wet), and EWL even after accounting for the highly significant phylogenetic signal of mass. T(b), BMR, C(wet), and EWL allometric residuals were correlated with some diet, distribution, and climatic variables before and after correction for phylogeny. T(b) residuals were higher for marsupials from arid environments (high T(a) and more variable rainfall). The fossorial marsupial mole had a lower-than-expected T(b) residual. The allometric slope for BMR was 0.72-0.75. Residuals were consistently related to distribution aridity and rainfall variability, with species from arid and variable rainfall habitats having a low BMR, presumably to conserve energy in a low-productivity environment. The nectarivorous honey possum had a higher-than-expected BMR. For C(wet), the allometric slope was 0.55-0.62; residuals were related to diet, with folivores having low and insectivores high C(wet) residuals. The allometric slope for EWL was 0.68-0.73. EWL residuals were consistently correlated with rainfall variability, presumably facilitating maintenance of water balance during dry periods.

Reexamining Echidna Physiology: The Big Picture for Tachyglossus aculeatus acanthion.

The early divergence of monotremes and therian mammals has resulted in considerable interest in the comparative physiology of the short-beaked echidna (Tachyglossus aculeatus), the most common and widespread living monotreme. However, there are many and varied interpretations of its physiology, reflecting the many and varied studies, limitations and uncertainties of aspects of some previous studies, and potential differences between the various subspecies. Consequently, we thoroughly examine here the standardized physiology of the most widely distributed subspecies of short-beaked echidna (T. aculeatus acanthion) over a wide range of ambient temperatures to definitively assess its physiology in a comparative context. We conclude that the low and variable body temperature of the short-beaked echidna is physiologically "primitive," but it also reflects adaptation to its myrmecophagous niche. Other aspects of its physiology are more typically mammalian. A low metabolic rate reflects its low body temperature, and ventilatory variables are matched to accommodate a modest gas exchange requirement. Thermal conductance is typical for a mammal of equivalent mass. In contrast to previous studies, we demonstrate that short-beaked echidnas can enhance evaporative water loss above thermoneutrality, like other mammals, with a similar capacity for evaporative heat loss. Cooling of their nasal blood sinus with nasal mucous may contribute to this enhanced evaporative cooling. Their capacity to evaporatively cool explains how their distribution can include habitats where ambient temperature, even in shelters, exceeds their supposed critical thermal limit.

A steady-state model of maximal oxygen and carbon dioxide transport in anuran amphibians.

A steady-state model, incorporating pulmonary ventilation, pulmonary diffusion capacity, cardiovascular transport capacity, and tissue diffusion capacity, was developed to describe the maximal O2 and CO2 transport capacity for an anuran amphibian (Bufo). Solution of the model by iterative calculation closely predicted 1) the empirical maximal O2 consumption (VO2max) for Bufo, 2) variation in empirical VO2max for three other genera (Rana, Xenopus, Scaphiopus), and the empirically observed effects on VO2max of 3) hypobaric hypoxia, 4) artificially induced anemia, and 5) beta-blockade of heart rate increment with activity. The model indicates that cardiovascular transport is the rate-limiting step to VO2max in amphibians and that an increase in circulatory O2 transport is a major physiological adaptation for increasing total aerobic capacity. CO2 transport and body fluid PCO2 values were primarily determined by pulmonary ventilatory capacity, and to a lesser extent by cardiovascular transport. The model should be generally applicable to other terrestrial vertebrates.

Dracunculiasis eradication: beginning of the end.

Beginning with the International Drinking Water Supply and Sanitation Decade (1981-1990), an increasingly broad coalition of international and bilateral agencies, organizations, private companies, and other institutions have joined forces to eradicate dracunculiasis (Guinea worm disease). From an estimated annual incidence of 10 million persons just before the campaign began, the remaining incidence of cases is now less than two million. More than 23,000 villages are known to be endemic. All 18 countries where the disease is still endemic have completed or begun nationwide searches to identify endemic villages, except Kenya. Dracunculiasis is nearly eradicated in Asia, where Pakistan found only 23 cases in 1992, and India found 1,081 cases. Cameroon and Senegal are close to achieving eradication in Africa, where the two formerly highest endemic countries, Nigeria and Ghana, reduced their combined total of cases from approximately 820,000 in 1989 to less than 240,000 in 1992. Much remains to be done, however, in francophone West Africa and especially in East Africa. The most serious current obstacles to eradicating dracunculiasis by 1995 are the civil war in Sudan, apathy of some national and international health officials, and inadequate funding for the campaign.

Dracunculiasis eradication: delayed, not denied.

By the end of 1998, Asia was free of dracunculiasis (Guinea worm disease), with Pakistan, India, and Yemen having interrupted transmission in 1993, 1996, and 1997, respectively. Transmission of the disease was also interrupted in Cameroon and Senegal during 1997. Chad reported only 3 cases during 1998. Dracunculiasis is now confined to only 13 countries in Africa. The overall number of cases has been reduced by more than 97% from the 3.2 million cases estimated to have occurred in 1986 to 78,557 cases reported in 1998. Because the civil war in Sudan remains the major impediment to eradication of dracunculiasis, the interim goal is to stop all transmission outside that country by the end of 2000. The most important operational need now is for national programs to improve the frequency and quality of supervision of village-based health workers in order to enhance the sensitivity of surveillance and effectiveness of case containment.

Dracunculiasis eradication: March 1994 update.

Substantial progress has been realized in the global campaign to eradicate dracunculiasis by the end of 1995 since a previous review of the subject was published in this journal a year ago. All known endemic countries are now engaged in the eradication effort, and one or more control measures are now in place in 93% of endemic villages. Despite improved surveillance for the disease, the number of reported cases of the disease has been reduced by 41% (to about 221,000), and the number of known endemic villages has been reduced by 28% (to about 16,500) in the past year. Priorities for national eradication programs in 1994 include increasing the use of vector control and intensifying the case containment strategy in endemic villages. It is still possible to achieve the eradication target of December 1995, but greatly intensified efforts this year will be required to do so.

Mobilization strategy for guinea worm eradication in Nigeria.

The transformation of dracunculiasis from an obscure and neglected rural disease to the highly visible target of a national eradication campaign in Nigeria is described in this report. This process progressed through four overlapping stages: documentation of the extent and nature of the disease as a national problem, demonstration in Nigeria that dracunculiasis could be effectively prevented by targeted provision and use of protected rural water supplies, mobilization for community participation in, and political support of, the eradication effort, and implementation of interventions nationwide. The conduct of the first national village-by-village search for cases and documentation of the adverse socioeconomic impact of the disease (e.g., on rice production) in Nigeria were the key elements used to solicit greater attention to the problem and mobilize support for its eradication. The critical role of the mass media in this effort and other benefits of this mobilization strategy are also highlighted.

Effects of urea on M4-lactate dehydrogenase from elasmobranchs and urea-accumulating Australian desert frogs.

We measured the effect of urea on M4-lactate dehydrogenase (M4-LDH) from elasmobranchs and Australian desert frogs (urea accumulators) and from two animals that do not accumulate urea, the axolotl and the rabbit. An analysis of the effect of urea on the Kd(NADH), V, V/K(m(prr)) and V/K(m(NADH)) shows that in all cases the major effect of urea was on the binding of pyruvate, which fits with data in the literature that show that urea acts as a competitive inhibitor of LDH. The characteristics of the elasmobranch enzymes are consistent with a proposed adaptation model, but the situation for the enzymes from the aestivating frogs is equivocal. Urea (400 mM) had less effect on the K(m(prr)) of M4-LDH from the urea accumulators than it did on the non-accumulators, suggesting a general adaptation and that the enzyme produced by the aestivating frogs (urea accumulators) is kinetically different from that of non-aestivating frogs (non-accumulators). A new approach is used to characterize the overall pattern of adaptation to urea. The pattern is similar in an enzyme from an elasmobranch and an aestivating frog despite the temporary presence of urea in the latter and the phylogenetic difference between these animals.

Termite digestibility and water and energy contents determine the water economy index of numbats (Myrmecobius fasciatus) and other myrmecophages.

Digestibility by captive numbats for termites was determined by feeding trials to be 81%+/-1.2% for Coptotermes sp. and 64%+/-3.3% for Nasutitermes sp. Water, ash, and energy content of both the Coptotermes (0.96+/-0.099 mg(dry mass) individual(-1), 78.0%+/-0.36% water, 5.8%+/-0.31% ash, 23.1+/-0.19 kJ g-1dry total energy) and Nasutitermes (0.91+/-0.046 mg(dry mass) individual(-1), 76.7%+/-3.09% water, 7.5%+/-1.10% ash, 22.7+/-0.36 kJ g-1dry total energy) were similar to values measured previously for other termites and for ants and insects in general. Numbats have a slow passage time for termites (20-30 h), presumably to enhance the digestion of termites. The water economy index (WEI) was 0.2 for captive numbats feeding on Coptotermes and 0.25 for Nasutitermes, whereas the WEI measured for wild, free-living numbats was 0.29, which corresponds to a digestibility of 58%. The WEI of a myrmecophage diet is determined by the energy and water contents and digestibilities of termites and ants, in the absence of drinking. The WEI for numbats, and other termitivorous mammals as well as reptiles, is higher than would be expected for an animal-based diet because of their relatively low digestibility (58%-81%) for termites. A high WEI preadapts myrmecophages to survival in arid environments without having to drink.

Effect of sampling regime on estimation of basal metabolic rate and standard evaporative water loss using flow-through respirometry.

Strict criteria have been established for measurement of basal metabolic rate and standard evaporative water loss to ensure that data can be compared intra- and interspecifically. However, data-sampling regimes vary, from essentially continuous sampling to interrupted (switching) systems with data recorded periodically at more widely spaced intervals. Here we compare one continuous and three interrupted sampling regimes to determine whether sampling regime has a significant effect on estimation of basal metabolic rate or standard evaporative water loss. Compared to continuous 20-s sampling averaged over 20 min, sampling every 6 min and averaging over 60 min overestimated basal metabolic rate and evaporative water loss, sampling every 3 min and averaging over 21 min underestimated basal metabolic rate, and sampling every 12 min and averaging over 36 min showed no difference in estimates. Increasing the period over which the minimum mean was calculated significantly increased estimates of physiological variables. Reducing the frequency of sampling from 20 s to a longer interval of 3, 6, or 12 min underestimated basal metabolic rate but not evaporative water loss. This indicates that sampling frequency per se influences estimates of basal metabolic rate and that differences are not just an artifact of differences in the period over which the mean is calculated. Sampling regime can have a highly significant influence on estimation of standard physiological variables, although the actual differences between sampling regimes were generally small (usually <5%). Although continuous sampling is the preferred sampling regime for open-flow respirometry studies, if time and cost are prohibitive, then use of an appropriate switching system will result in smaller errors than measuring individuals continuously for shorter periods.

Effects of measurement duration on the determination of basal metabolic rate and evaporative water loss of small marsupials: how long is long enough?

We examined the time course for measurement of basal metabolic rate (BMR; measured as O(2) consumption and CO(2) production) and standard evaporative water loss (EWL) for six species of small marsupial to determine the minimum time required to achieve basal/standard values. There was a highly significant effect of measurement duration on measured physiological variables with values for O(2) consumption, CO(2) production, and EWL decreasing with time for all species. The time required to attain values statistically indistinguishable from minimal differed significantly between species, but in general O(2) consumption rate reached basal values after 4.3 h, CO(2) production after 4.5 h, and evaporative water loss after 5.2 h. For 16 BMR measurements of small marsupial species in the literature, with experimental duration provided, 10 were for less than 4 h, suggesting that their BMR values might be overestimates. For EWL, three of the four published values for small marsupials may be overestimates. It is clear that appropriate experimental duration is an important component of the measurement protocol for both BMR and standardized water loss, which needs to be rigorously observed in future studies.

Metabolic, ventilatory, and hygric physiology of the gracile mouse opossum (Gracilinanus agilis).

We present the first complete study of basic laboratory-measured physiological variables (metabolism, thermoregulation, evaporative water loss, and ventilation) for a South American marsupial, the gracile mouse opossum (Gracilinanus agilis). Body temperature (T(b)) was thermolabile below thermoneutrality (T(b) = 33.5 degrees C), but a substantial gradient between T(b) and ambient temperature (T(a)) was sustained even at T(a) = 12 degrees C (T(b) = 30.6 degrees C). Basal metabolic rate of 1.00 mL O2 g(-1) h(-1) at T(a) = 30 degrees C conformed to the general allometric relationship for marsupials, as did wet thermal conductance (5.7 mL O2 g(-1) h(-1) degrees C(-1)). Respiratory rate, tidal volume, and minute volume at thermoneutrality matched metabolic demand such that O2 extraction was 12.4%, and ventilation increased in proportion to metabolic rate at low T(a). Ventilatory accommodation of increased metabolic rate at low T(a) was by an increase in respiratory rate rather than by tidal volume or O2 extraction. Evaporative water loss at the lower limit of thermoneutrality conformed to that of other marsupials. Relative water economy was negative at thermoneutrality but positive below T(a) = 12 degrees C. Interestingly, the Neotropical gracile mouse opossums have a more positive water economy at low T(a) than an Australian arid-zone marsupial, perhaps reflecting seasonal variation in water availability for the mouse opossum. Torpor occurred at low T(a), with spontaneous arousal when T(b) > 20 degrees C. Torpor resulted in absolute energy and water savings but lower relative water economy. We found no evidence that gracile mouse opossums differ physiologically from other marsupials, despite their Neotropical distribution, sympatry with placental mammals, and long period of separation from Australian marsupials.

Allometry of evaporative water loss in marsupials: implications of the effect of ambient relative humidity on the physiology of brushtail possums (Trichosurus vulpecula).

To better understand the effects of ambient relative humidity (RH) on physiological variables and the implications of RH-correcting evaporative water loss (EWL) data for marsupials, we examined the effect of RH on EWL, body temperature (Tb), metabolic rate (MR) and thermal conductance (C) of the brushtail possum (Trichosurus vulpecula), a medium-sized marsupial. Correcting EWL data for 27 species of marsupial for water vapour pressure deficit (DeltaWVP) in the chamber during measurement significantly increased, rather than decreased, the variability of the allometric relationship for EWL. For the brushtail possum, both ambient temperature (Ta) and RH significantly affected EWL. At Ta=25 degrees C, EWL was independent of RH at "63% RH, but decreased linearly at higher RH values. At Ta=30 degrees C, EWL was significantly related to RH from 26% to 92% RH. There was a significant effect of Ta on Tb and dry thermal conductance (Cdry; higher at 30 degrees C), but no effect of RH. For MR and wet thermal conductance (Cwet) there was a significant effect of Ta (MR higher and Cwet lower at 25 degrees C), and RH at Ta=30 degrees C (MR higher and Cwet lower at the lowest RH) but not at 25 degrees C. Our results indicate that brushtail possums do not necessarily show the linear relationship between ambient RH and EWL expected for an endotherm, possibly because of behavioural modification of their immediate microclimate. This may account for the failure of WVP deficit correction to improve the allometric EWL relationship for marsupials. Chamber RH is an important environmental factor to be considered when measuring standard physiological variables such as MR and Cwet.

Numbats and aardwolves--how low is low? A re-affirmation of the need for statistical rigour in evaluating regression predictions.

Many comparative physiological studies aim to determine if a particular species differs from a prediction based on a linear allometric regression for other species. However, the judgment as to whether the species in question conforms to this allometric relationship is often not based on any formal statistical analysis. An appropriate statistical method is to compare the new species' value with the 95% confidence limits for predicting an additional datum from the relationship for the other species. We examine the basal metabolic rate (BMR) of the termitivorous numbat (Myrmecobius fasciatus) and aardwolf (Proteles cristatus) to demonstrate the use of the 95% prediction limits to determine statistically if they have a lower-than-expected BMR compared to related species. The numbat's BMR was 83.6% of expected from mass, but fell inside the 95% prediction limits for a further datum; a BMR < 72.5% of predicted was required to fall below the one-tail 95% prediction limits. The aardwolf had a BMR that was only 74.2% of predicted from the allometric equation, but it also fell well within the 95% prediction limits; a BMR of only 41.8% of predicted was necessary to fall below the one-tail 95% prediction limits. We conclude that a formal statistical approach is essential, although it is difficult to demonstrate that a single species statistically differs from a regression relationship for other species.

Water relations of the burrowing sandhill frog, Arenophryne rotunda (Myobatrachidae).

Arenophryne rotunda is a small (2-8 g) terrestrial frog that inhabits the coastal sand dunes of central Western Australia. While sand burrowing is a strategy employed by many frog species inhabiting Australia's semi-arid and arid zones, A. rotunda is unique among burrowing species because it lives independently of free water and can be found nocturnally active on the dune surface for relatively extended periods. Consequently, we examined the physiological factors that enable this unique frog to maintain water balance. A. rotunda was not found to have any special adaptation to reduce EWL (being equivalent to a free water surface) or rehydrate from water (having the lowest rehydration rate measured for 15 Western Australian frog species), but it was able to maintain water balance in sand of very low moisture (1-2%). Frogs excavated in the field were in dune sand of 4.4% moisture content, as a consequence of recent rain, which was more than adequate for these frogs to maintain water balance as reflected by their low plasma and urine osmotic concentrations. We suggest that in dry periods of the year, A. rotunda can achieve positive water balance by cutaneous water uptake by burrowing deeper into the substrate to where the percent water content is greater than 1.5%.

Significant decrease in the prevalence of Wuchereria bancrofti infection in anopheline mosquitoes following the addition of albendazole to annual, ivermectin-based, mass treatments in Nigeria.

A prospective entomological survey was conducted in four sentinel villages in central Nigeria from 1999-2002, to assess the impact of annual, single-dose, mass drug administrations (MDA), with a combination of ivermectin and albendazole, on the transmission of Wuchereria bancrofti. As they were also endemic for human onchocerciasis, the four villages had received annual MDA based on ivermectin alone for 7 years prior to the addition of albendazole. Resting Anophelines gambiae s. l., An. funestus and Culex species were collected from 92 sequentially sampled households and dissected. Mosquitoes harbouring any larval stage of W. bancrofti were classified as 'infected', and those containing the third-stage larvae of the parasite were classified as 'infective'. Over the 41-month observation period, 4407 mosquitoes were captured and dissected, of which 64% were An. gambiae s. l., 34% An. funestus, and 1% Culex species. The baseline data, from dissections performed before the addition of albendazole to the MDA, showed high prevalences of mosquito infection (8.9%) and infectivity (2.9%), despite apparently good treatment coverages during the years of annual ivermectin monotherapy. Only the anopheline mosquitoes were found to harbour W. bancrofti larvae. After the third round of MDA with the ivermectin-albendazole combination, statistically significant decreases in the prevalences of mosquito infection (down to 0.6%) and infectivity (down to 0.4%) were observed (P<0.0001 for each). The combination of albendazole and ivermectin appears to be superior to ivermectin alone for reducing the frequency of W. bancrofti infection in mosquitoes.

Metabolic, respiratory and haematological adjustments of the little pocket mouse to circadian torpor cycles.

Metabolic, respiratory and haematological parameters were investigated for the Little Pocket mouse during circadian torpor cycles. The rate of O2 consumption decreased from 7.04 to 0.05 ml O2.g-1.hr-1, with a corresponding decrease in respiratory minute volume from 49.4 to 0.9 ml.min-1 during torpor at an ambient temperature of 10 C. No changes in haemoglobin concentration (19.7 g/100 ml), haematocrit (54%), red blood corpuscle count (12.4 10(6)/microliter), mean corpuscular volume (43.6 micrometer3), mean corpuscular haemoglobin content (16.2 pg), mean corpuscular haemoglobin concentration (37.4%) and [2,3-DPG] (9.6 mumol/g Hb) were observed during torpor cycles. The half saturation tension of P. longimembris haemoglobin was 41 mm Hg (37 C, pH = 7.28) and 19.7 mm Hg (10 degrees C, pH = 7.51). The effect of temperature on P50 was deltalog P50/ C = +0.0106 (pH = 7.4). Venous blood parameters were: euthermic mice (37 C); PCO2 = 36.8 mm Hg, PO2 = 49.5 mm Hg, pH = 7.28, [HCO-3] = 17.3 mmol/l; torpid mice (10 C); PCO2 = 14.6, PO2 = 35.7 pH = 7.51, [HCO-3] = 18.8. These data indicate a new, relatively acidotic acid-base status during torpor, characterised by a higher H+/ OH- ratio. The respiratory sensitivity to inspired CO2 of pocket mice was, despite their being semi-fossorial, typical of other mammals. High concentrations of CO2 did not induce, or facilitate, entry into torpor.