Daily torpor and energetics in a tropical mammal, the northern blossom-bat Macroglossus minimus (Megachiroptera)

Little is known about torpor in the tropics or torpor in megachiropteran species. We investigated thermoregulation, energetics and patterns of torpor in the northern blossom-bat Macroglossus minimus (16 g) to test whether physiological variables may explain why its range is limited to tropical regions. Normothermic bats showed a large variation in body temperature (T _b) (33 to 37 °C) over a wide range of ambient temperatures (T _as) and a relatively low basal metabolic rate (1.29 ml O₂ g⁻¹h⁻¹). Bats entered torpor frequently in the laboratory at T _as between 14 and 25 °C. Entry into torpor always occurred when lights were switched on in the morning, independent of T _a. MRs during torpor were reduced to about 20–40% of normothermic bats and T _bs were regulated at a minimum of 23.1 ± 1.4 °C. The duration of torpor bouts increased with decreasing T _a in non-thermoregulating bats, but generally terminated after 8 h in thermoregulating torpid bats. Both the mean minimum T _b and MR of torpid M. minimus were higher than that predicted for a 16-g daily heterotherm and the T _b was also about 5 °C higher than that of the common blossom-bat Syconycteris australis, which has a more subtropical distribution. These observations suggest that variables associated with torpor are affected by T _a and that the restriction to tropical areas in M. minimus to some extent may be due to their ability to enter only very shallow daily torpor. Accepted: 22 September 1997     

Torpor, thermal biology, and energetics in Australian long-eared bats (Nyctophilus)

Previous studies have suggested that Australian long-eared bats (Nyctophilus) differ from northern-hemisphere bats with respect to their thermal physiology and patterns of torpor. To determine whether this is a general trait of Australian bats, we characterised the temporal organisation of torpor and quantified metabolic rates and body temperatures of normothermic and torpid Australian bats (Nyctophilus geoffroyi, 7 g and N. gouldi, 10 g) over a range of air temperatures and in different seasons. The basal metabolic rate of normothermic bats was 1.36 ± 0.17 ml g⁻¹ h⁻¹ (N. geoffroyi) and 1.22 ± 0.13 ml g⁻¹ h⁻¹ (N. gouldi), about 65% of that predicted by allometric equations, and the corresponding body temperature was about 36 °C. Below an air temperature of about 25 °C bats usually remained normothermic for only brief periods and typically entered torpor. Arousal from torpor usually occurred shortly after the beginning of the dark phase and torpor re-entry occurred almost always during the dark phase after normothermic periods of only 111 ± 48 min (N. geoffroyi) and 115 ± 66 min (N. gouldi). At air temperatures below 10 °C, bats remained torpid for more than 1 day. Bats that were measured overnight had steady-state torpor metabolic rates representing only 2.7% (N. geoffroyi) and 4.2% (N. gouldi) of the basal metabolic rate, and their body temperatures fell to minima of 1.4 and 2.3 °C, respectively. In contrast, bats measured entirely during the day, as in previous studies, had torpor metabolic rates that were up to ten times higher than those measured overnight. The steady-state torpor metabolic rate of thermoconforming torpid bats showed an exponential relationship with body temperature (r ² = 0.94), suggesting that temperature effects are important for reduction of metabolic rate below basal levels. However, the 75% reduction of metabolic rate between basal metabolic rate and torpor metabolic rate at a body temperature of 29.3 °C suggests that metabolic inhibition also plays an important role. Torpor metabolic rate showed little or no seasonal change. Our study suggests that Australian Nyctophilus bats have a low basal metabolic rate and that their patterns of torpor are similar to those measured in bats from the northern hemisphere. The low basal metabolic rate and the high proclivity of these bats for using torpor suggest that they are constrained by limited energy availability and that heterothermy plays a key role in their natural biology. Accepted: 22 November 1999     

Comparison of hibernation, estivation and daily torpor in the edible dormouse, Glis glis

Three major forms of dormancy in mammals have been classified: hibernation in endotherms is characterised by reduced metabolic rate (MR) and body temperature (T _b) near ambient temperature (T _a) over prolonged times in the winter. Estivation is a similar form of dormancy in a dry and hot environment during summertime. Daily torpor is defined as reduced MR and T _b lower than 32 °C, limited to a duration of less than 24 h. The edible dormouse (Glis glis) is capable for all three distinct forms of dormancy. During periods of food restriction and/or low T _a, daily torpor is displayed throughout the year, alternating with hibernation and estivation in winter and summer respectively. We recorded T _b, O₂-consumption and CO₂-production in unrestrained dormice at different T _a's for periods of up to several months. Cooling rate and rate of metabolic depression during entrance into the torpid state was identical in all three forms of dormancy. The same was true for thermal conductance, maximum heat production, duration of arousal and cost of an arousal. The only difference between hibernation and daily torpor was found in the bout duration. A daily torpor bout lasted 3–21 h, a hibernation bout 39–768 h. As a consequence of prolonged duration, MR, T _b and also the T _b − T _a gradient decreased to lower values during hibernation bouts when compared to daily torpor bouts. Our findings suggest that all three forms of dormancy are based on the same physiological mechanism of thermal and metabolic regulation. Accepted: 27 June 2000     

Dietary cholesterol enhances torpor in a rodent hibernator

Dietary cholesterol can affect both body lipid composition and steroid hormone concentration. We investigated whether a diet rich in cholesterol influences torpor patterns of hibernating chipmunks (Tamias amoenus) and, if so, whether these changes are better explained by diet-induced changes in body lipid composition or the concentration of testosterone, which at high levels inhibits torpor. Two groups of chipmunks were maintained either on a cholesterol diet (rodent chow containing 10% cholesterol) or a control diet (rodent chow) during pre-hibernation fattening and throughout the hibernation season. Torpid chipmunks on the cholesterol diet had significantly lower minimum body temperatures (−0.2 ± 0.2 vs +0.6 ± 0.2 °C), lower metabolic rates (0.029 ± 0.002 ml O₂ g⁻¹h⁻¹ vs 0.035 ± 0.001 ml O₂ g⁻¹h⁻¹), and longer torpor bouts at −1 °C (6.8 ± 0.5 vs 4.1 ± 1.0 days) than chipmunks on the control diet. Dietary cholesterol resulted in a significant increase in blood plasma cholesterol (sevenfold), liver cholesterol content (6.9-fold) and liver triglyceride content (3.5-fold) in comparison to controls. In contrast, dietary cholesterol had no detectable effect on the concentration of plasma testosterone, which was very low in both groups. Since torpor was deeper and longer in animals on the cholesterol diet our study suggests that torpor patterns of chipmunks were either directly affected by the dietary cholesterol or via changes in body lipid composition. Accepted: 22 January 1997     

Daily energy expenditure of the grey mouse lemur (Microcebus murinus): a small primate that uses torpor

We aimed to investigate the pattern of utilisation of torpor and its impact on energy budgets in free-living grey mouse lemurs (Microcebus murinus), a small nocturnal primate endemic to Madagascar. We measured daily energy expenditure (DEE) and water turnover using doubly labelled water, and we used temperature-sensitive radio collars to measure skin temperature (T _sk) and home range. Our results showed that male and female mouse lemurs in the wild enter torpor spontaneously over a wide range of ambient temperatures (T _a) during the dry season, but not during the rainy season. Mouse lemurs remained torpid between 1.7–8.9 h with a daily mean of 3.4 h, and their T _sk s fell to a minimum of 18.8 °C. Mean home ranges of mouse lemurs which remained normothermic were similar in the rainy and dry season. During the dry season, the mean home range of mouse lemurs showing daily torpor was significantly smaller than that of animals remaining normothermic. The DEE of M. murinus remaining normothermic in the rainy season (122 ± 65.4 kJ day⁻¹) was about the same of that of normothermic mouse lemurs in the dry season (115.5 ± 27.3 kJ day⁻¹). During the dry season, the mean DEE of M. murinus that utilised daily torpor was 103.4 ± 32.7 kJ day⁻¹ which is not significantly different from the mean DEE of animals remaining normothermic. We found that the DEE of mouse lemurs using daily torpor was significantly correlated with the mean temperature difference between T _sk and T _a (r ²=0.37) and with torpor bout length (r ² =0.46), while none of these factors explained significant amounts of variation in the DEE of the mouse lemurs remaining normothermic. The mean water flux rate of mouse lemurs using daily torpor (13.0 ± 4.1 ml day⁻¹) was significantly lower than that of mouse lemurs remaining normothermic (19.4 ± 3.8 ml day⁻¹), suggesting the lemurs conserve water by entering torpor. Thus, this first study on the energy budget of free-ranging M. murinus demonstrates that torpor may not only reflect its impact on the daily energy demands, but involve wider adaptive implications such as water requirements. Accepted: 29 August 2000     

Torpor and thermal energetics in a tiny Australian vespertilionid,the little forest bat (<Emphasis Type="Italic">Vespadelus vulturnus</Emphasis>)

Data on thermal energetics for vespertilionid bats are under-represented in the literature relative to their abundance, as are data for bats of very small body mass. Therefore, we studied torpor use and thermal energetics in one of the smallest (4 g) Australian vespertilionids, Vespadelus vulturnus. We used open-flow respirometry to quantify temporal patterns of torpor use, upper and lower critical temperatures (T _uc and T _lc) of the thermoneutral zone (TNZ), basal metabolic rate (BMR), resting metabolic rate (RMR), torpid metabolic rate (TMR), and wet thermal conductance (C _wet) over a range of ambient temperatures (T _a). We also measured body temperature (T _b) during torpor and normothermia. Bats showed a high proclivity for torpor and typically aroused only for brief periods. The TNZ ranged from 27.6°C to 33.3°C. Within the TNZ T _b was 33.3±0.4°C and BMR was 1.02±0.29 mlO₂ g⁻¹ h⁻¹ (5.60±1.65 mW g⁻¹) at a mean body mass of 4.0±0.69 g, which is 55 % of that predicted for a 4 g bat. Minimum TMR of torpid bats was 0.014±0.006 mlO₂ g⁻¹ h⁻¹ (0.079±0.032 mW g⁻¹) at T _a=4.6±0.4°C and T _b=7.5±1.9. T _lc and C _wet of normothermic bats were both lower than that predicted for a 4 g bat, which indicates that V. vulturnus is adapted to minimising heat loss at low T _a. Our findings support the hypothesis that vespertilionid bats have evolved energy-conserving physiological traits, such as low BMR and proclivity for torpor.     

Field metabolic rates and water uptake in the blossom-bat Syconycteris australis (Megachiroptera)

Blossom-bats, Syconycteris australis (18 g) are known to be highly active throughout the night. Since this species frequently enters torpor, we postulated that their use of heterothermy may be related to a high energy expenditure in the field. To test this hypothesis we measured field metabolic rates (FMR) of S. australis at a subtropical site using the doubly labelled water (DLW) method. We also measured DLW turnover in captive animals held at constant ambient temperature (T _a) with ad libitum food to estimate whether T _a and food availability affect energy expenditure under natural conditions. The FMR of S. australis was 8.55 ml CO₂ g⁻¹h⁻¹ or 76.87 kJ day⁻¹ which is 7.04 times the basal metabolic rate (BMR) and one of the highest values reported for endotherms to date. Mass-specific energy expenditure by bats in the laboratory was about two-thirds of that of bats in the field, but some of this difference was explained by the greater body mass in captive bats. This suggests that foraging times in the field and laboratory were similar, and daily energy expenditure was not strongly affected by T _a or ad libitum food. Water uptake in the field was significantly higher than in the laboratory, most likely because nectar contained more water than the laboratory diet. Our study shows that S. australis has a FMR that is about double that predicted for its size although its BMR is lower than predicted. This supports the view that caution must be used in making assumptions from measurements of BMR in the laboratory about energy and other biological requirements in free-ranging animals. Accepted: 4 January 1999     

Hibernation by a free-ranging subtropical bat (Nyctophilus bifax)

Knowledge about torpor in free-ranging subtropical bats is scarce and it is widely believed that low and stable ambient temperatures are necessary for prolonged torpor. We present temperature-telemetry data from free-ranging male (n = 4) and female (n = 4) subtropical vespertilionid bats, Nyctophilus bifax (~10 g), exposed to pronounced daily fluctuations of ambient temperature. All bats used torpor on every day in winter and both males and females exhibited multi-day torpor bouts of up to 5.4 days. Although females were larger than males, patterns of torpor were similar in both sexes. Torpor use was correlated with prevailing weather conditions and, on days when bats remained torpid, maximum ambient temperature was significantly lower than on days when bats aroused. Moreover, the duration of interbout normothermic periods at night increased with increasing average nightly ambient temperature. Skin temperature of torpid bats varied by 10.2 ± 3.6°C day⁻¹ (n = 8, N = 47) and daily minimum skin temperature was positively correlated with the daily minimum ambient temperature. Our study shows that prolonged torpor is an important component of the winter ecology of a subtropical bat and that torpor and activity patterns of N. bifax predominantly reflect prevailing weather conditions.     

Does fear beget fear? Risk-mediated habitat selection triggers predator avoidance at lower trophic levels

Seasonal changes in weather and food availability differentially impact energy budgets of small mammals such as bats. While most thermal physiological research has focused on species that experience extreme seasonal temperature variations, knowledge is lacking from less variable temperate to subtropical climates. We quantified ambient temperature (T _a) and skin temperature (T _sk) responses by individuals from a population of New Zealand lesser short-tailed bats (Mystacina tuberculata) during summer and winter using temperature telemetry. During summer, communal roosts were more thermally stable than T _a. During winter, solitary roosts were warmer than T _a indicating significant thermal buffering. Communal roost trees were used on 83 % of observation days during summer, and individuals occupying them rarely entered torpor. Solitary roosts were occupied on 93 % of observation days during winter, and 100 % of individuals occupying them used torpor. During summer and winter, bats employed torpor on 11 and 95 % of observation days, respectively. Maximum torpor bout duration was 120.8 h and winter torpor bout duration correlated negatively with mean T _a. Torpor bout duration did not differ between sexes, although female minimum T _sk was significantly lower than males. The summer Heterothermy Index varied, and was also significantly affected by T _a. Mean arousal time was correlated with sunset time and arousals occurred most frequently on significantly warmer evenings, which are likely associated with an increased probability of foraging success. We provide the first evidence that torpor is used flexibly throughout the year by M. tuberculata, demonstrating that roost choice and season impact torpor patterns. Our results add to the growing knowledge that even small changes in seasonal climate can have large effects on the energy balance of small mammals.     

Water and sodium balances and metabolic physiology of house mice (Mus domesticus) and short-tailed mice (Leggadina lakedownensis) under laboratory conditions

A laboratory study investigated the metabolic physiology, and response to variable periods of water and sodium supply, of two arid-zone rodents, the house mouse (Mus domesticus) and the Lakeland Downs short-tailed mouse (Leggadina lakedownensis) under controlled conditions. Fractional water fluxes for M. domesticus (24 ± 0.8%) were significantly higher than those of L. lakedownensis (17 ± 0.7%) when provided with food ad libitum. In addition, the amount of water produced by M. domesticus and by L. lakedownensis from metabolic processes (1.3 ± 0.4 ml · day⁻¹ and 1.2 ± 0.4 ml · day⁻¹, respectively) was insufficient to provide them with their minimum water requirement (1.4 ± 0.2 ml · day⁻¹ and 2.0 ± 0.3 ml · day⁻¹, respectively). For both species of rodent, evaporative water loss was lowest at 25 °C, but remained significantly higher in M. domesticus (1.1 ± 0.1 mg H₂O · g^−0.122 · h⁻¹) than in L. lakedownensis (0.6 ± 0.1 mg H₂O · g^−0.122 · h⁻¹). When deprived of drinking water, mice of both species initially lost body mass, but regained it within 18 days following an increase in the amount of seed consumed. Both species were capable of drinking water of variable saline concentrations up to 1 mol · l⁻¹, and compensated for the increased sodium in the water by excreting more urine to remove the sodium. Basal metabolic rate was significantly higher in M. domesticus (3.3 ± 0.2 mg O₂ · g^−0.75 · h⁻¹) than in L. lakedownensis (2.5 ± 0.1 mg O₂ · g^−0.75 · h⁻¹). The study provides good evidence that water flux differences between M. domesticus and L. lakedownensis in the field are due to a requirement for more water in M. domesticus to meet their physiological and metabolic demands. Sodium fluxes were lower than those observed in free-ranging mice, whose relatively high sodium fluxes may reflect sodium associated with available food. Accepted: 16 August 1999     

Energetic cost of hovering flight in nectar-feeding bats (Phyllostomidae: Glossophaginae) and its scaling in moths, birds and bats

Three groups of specialist nectar-feeders covering a continuous size range from insects, birds and bats have evolved the ability for hovering flight. Among birds and bats these groups generally comprise small species, suggesting a relationship between hovering ability and size. In this study we established the scaling relationship of hovering power with body mass for nectar-feeding glossophagine bats (Phyllostomidae). Employing both standard and fast-response respirometry, we determined rates of gas exchange in Hylonycteris underwoodi (7 g) and Choeronycteris mexicana (13–18 g) during hover-feeding flights at an artificial flower that served as a respirometric mask to estimate metabolic power input. The O₂ uptake rate (V˙ _o2) in ml g⁻¹ h⁻¹ (and derived power input) was 27.3 (1.12 W or 160 W kg⁻¹) in 7-g Hylonycteris and 27.3 (2.63 W or 160 W kg⁻¹) in 16.5-g Choeronycteris and thus consistent with measurements in 11.9-g Glossophagasoricina (158 W kg⁻¹, Winter 1998). V˙ _o2 at the onset of hovering was also used to estimate power during forward flight, because after a transition from level forward to hovering flight gas exchange rates initially still reflect forward flight rates. V˙ _o2 during short hovering events (<1.5 s) was 19.0 ml g⁻¹ h⁻¹ (1.8 W) in 16-g Choeronycteris, which was not significantly different from a previous, indirect estimate of the cost of level forward flight (2.1 W, Winter and von Helversen 1998). Our estimates suggest that power input during hovering flight P _h (W) increased with body mass M (kg) within 13–18-g Choeronycteris (n = 4) as P _h  = 3544 (±2057 SE) M ^{1.76 (±0.21 SE)} and between different glossophagine bat species (n = 3) as P _h  = 128 (±2.4 SE) M ^{0.95 (±0.034 SE)}. The slopes of three scaling functions for flight power (hovering, level forward flight at intermediate speed and submaximal flight power) indicate that: 1. The relationship between flight power to flight speed may change with body mass in the 6–30-g bats from a J- towards a U-shaped curve. 2. A metabolic constraint (hovering flight power equal maximal flight power) may influence the upper size limit of 30–35 g for this group of flower specialists. Mass-specific power input (W kg⁻¹) during hovering flight appeared constant with regard to body size (for the mass ranges considered), but differed significantly (P < 0.001) between groups. Group means were 393 W kg⁻¹ (sphingid moths), 261 W kg⁻¹ (hummingbirds) and 159 W kg⁻¹ (glossophagine bats). Thus, glossophagine bats expend the least metabolic power per unit of body mass supported during hovering flight. At a metabolic power input of 1.1 W a glossophagine bat can generate the lift forces necessary for balancing 7 g against gravitation, whereas a hummingbird can support 4 g and a sphingid moth only 3 g of body mass with the same amount of metabolic energy. These differences in power input were not fully explained by differences in induced power output estimated from Rankine-Froude momentum-jet theory. Accepted: 10 November 1998     

Metabolic rates in an anadromous clupeid, the American shad (Alosa sapidissima)

To assess the energetics of migration in an anadromous fish, adult American shad (Alosa sapidissima) were swum in a large respirometer at a range of speeds (1.0–2.3 body lengths (BL) s⁻¹, 13–24 °C). Metabolic rate (M_O2) was logarithmically related to swimming speed (Bl s⁻¹; r ² = 0.41, slope = 0.23 ± 0.037) and tailbeat frequency (beats × min⁻¹; r ² = 0.52, slope = 0.003 ± 0.0003). Temperature had a significant effect on metabolic rate (r ² = 0.41) with a Q₁₀ of 2.2. Standard metabolic rate (SMR), determined directly after immobilization with the neuroblocker gallamine triethiodide, ranged from 2.2–6.2 mmolO₂ kg⁻¹ h⁻¹ and scaled with mass (W) such that SMR = 4.0 (±0.03)W^{0.695(±0.15)}. Comparison of directly determined and extrapolated SMR suggests that swimming respirometry provides a good estimate of SMR in this species, given the differences in basal activity monitored by the two methods. Overall, American shad metabolic rates (M_O2 and SMR) were intermediate between salmonids and fast-swimming perciforms, including tunas, and may be a result of evolutionary adaptation to their active pelagic, schooling life history. This study demonstrates variability in metabolic strategy among anadromous fishes that may be important to understanding the relative success of different migratory species under varying environmental conditions. Accepted: 3 March 1999     

Exchanges of oxygen, carbon dioxide, nitrogen and water in the caecilian Dermophis mexicanus

Oxygen consumption was measured in five Dermophis mexicanus and averaged (±SEM) 0.047 ± 0.004 ml O₂ g⁻¹ h⁻¹. Carbon dioxide production averaged 0.053 ± 0.005 ml CO₂ g⁻¹ h⁻¹ in the same five animals 1 week later. This metabolic rate is similar to metabolic rates of other Gymnophionans but lower than metabolic rates reported for Anurans and Urodeles. Total nitrogen excretion averaged 1.37 μmol N g⁻¹ h⁻¹ which is higher than that found for other amphibians. Of this, 82.5% (1.13 μmol N g⁻¹ h⁻¹) was in the form of urea while 17.5% (0.24 μmol N g⁻¹ h⁻¹) was in the form of NH₃ + NH⁺ ₄. Such ureotelism is typical of terrestrial amphibians like D. mexicanus. Osmotic water flux averaged 0.0193 ml g⁻¹ h⁻¹ in control (sham injected) animals and was not significantly altered by injection of either arginine vasotocin or mesotocin. This osmotic flux is similar to osmotic fluxes found for other terrestrial amphibians. The combined data suggest that metabolism in D. mexicanus is, like most other Gymnophionans, lower than other amphibians. The high rates of nitrogen (especially urea) excretion suggests that this fossorial animal accumulates urea like other burrowing amphibians. Accepted: 27 June 2000     

Energetics during nursing and early postweaning fasting in hooded seal (Cystophora cristata ) pups from the Gulf of St Lawrence, Canada

In this study we measured growth and milk intake and calculated energy intake and its allocation into metabolism and stored tissue for hooded seal (Cystophora cristata) pups. In addition, we measured mass loss, change in body composition and metabolic rate during the first days of the postweaning fast. The mean body mass of the hooded seal pups (n = 5) at the start of the experiments, when they were new-born, was 24.3 ± 1.3 kg (SD). They gained an average of 5.9 ± 1.1. kg · day⁻¹ of which 19% was water, 76% fat and 5% protein. This corresponds to an average daily energy deposition of 179.8 ± 16.0 MJ. The pups were weaned at an average body mass of 42.5 ± 1.0 kg 3.1 days after the experiment was initiated. During the first days of the postweaning fast the pups lost an average of 1.3 ± 0.5␣kg of body mass daily, of which 56% was water, 16% fat and 28% protein. During the nursing period the average daily water influx for the pups was 124.6 ± 25.8 ml · kg⁻¹. The average CO₂ production during this period was 1.10 ± 0.20 ml · g⁻¹ · h⁻¹, which corresponds to a field metabolic rate of 714 ± 130 kJ ·  kg⁻¹ · day⁻¹, or 5.8 ± 1.1 times the predicted basal metabolic rate according to Kleiber (1975). During the postweaning fast the average daily water influx was reduced to 16.1 ± 6.6 ml · kg⁻¹. The average CO₂ production in␣this period was 0.58 ± 0.17 ml · g⁻¹ · h⁻¹ which corresponds to a field metabolic rate of 375 ± 108 kJ · kg⁻¹ · day⁻¹ or 3.2 ± 0.9 times the predicted basal metabolic rate. Average values for milk composition were 33.5% water, 58.6% fat and 6.2% protein. The pups drank an average of 10.4 ± 1.8␣kg of milk daily, which represents an energy intake of 248.9 ± 39.1 MJ · day⁻¹. The pups were able to store 73.2 ± 7.7% of this energy as body tissue. Accepted: 15 August 1996     

Hypoxia tolerance associated with activity reduction is a key adaptation for Laternula elliptica seasonal energetics

Seasonal dormancy is a widespread mechanism for reducing energy expenditure during periods of low energy availability. Seasonal variation in activity and the cost of pumping water through the siphons were investigated to estimate the importance of activity regulation to the seasonal energy budget of the Antarctic clam, Laternula elliptica. In the laboratory, a metabolic rate of 26.35 μmol O₂ h⁻¹ was estimated for a 50-mm shell length L. elliptica pumping water at −0.4 °C. In the field, the proportion of time siphons were visible at the sediment surface varied seasonally (32% visible in June/July compared to 86% in December/January). L. elliptica were actively pumping for a minimum of 19% of each 24-h period during winter (August) compared to a summer maximum when animals were actively pumping for 73% of the time (February). This resulted in a 3.7-fold seasonal difference in the calculated energy consumption of a 50-mm L. elliptica (19.2 μmol O₂ h⁻¹ in February versus 5.0 μmol O₂ h⁻¹ in August), which closely matches the 3.0-fold seasonal variation in metabolic rate found previously. Seasonal variation in activity could therefore be responsible for much of the seasonal difference in energy consumption of L. elliptica. Inter-annual variation in timing of the seasonal activity maxima (January 2004 and March 1999) was correlated with variation in the timing of the summer plankton bloom in Ryder Bay. In the laboratory, periods of extended siphon closure (133 ± 114 min, mean ± SD) were accompanied by long periods of heart arrhythmia (167 ± 135 min), during which time blood oxygen levels dropped to values close to zero. Heart arrhythmia is most likely part of a hypo-metabolic adaptation to reduce energy costs during extended periods of siphon closure. Physiological and behavioural dormancy, with associated hypoxia tolerance, appear to be key mechanisms controlling the seasonal energy budget of L. elliptica.     

A neutral carrier-based liquid membrane microelectrode for divalent putrescine cations

A new ion-selective liquid membrane microelectrode, based on the neutral carrier 1,1′-bis(2,3-naphtho-18-crown-6), is described that shows the dependence of EMF on the activity of divalent putrescine cations a _Put, with the linear slope s _Put = 26 ± 3 mV/decade (mean ± SD, N = 18), in the range 10⁻⁴–10⁻¹ M at 25 ± 1 °C. Values of potentiometric putrescine cation selectivity coefficients of logK ^Pot _{Put j} (mean ± SD, N) are obtained by the separate solution method for the ions K⁺ (1.0 ± 0.4, 10), Na⁺ (−1.2 ± 0.4, 8), Ca²⁺ (−2.3 ± 0.5, 10) and Mg²⁺ (−2.5 ± 0.5, 7). The microelectrode can be applied for the direct analysis of the activities of free divalent putrescine cations in the range 5 × 10⁻⁴ to 10⁻¹ M in an extracellular ionic environment. Established analytical methods, e.g. high performance liquid chromatography, determine the total concentration of the derivatives of free and bound putrescine. Received: 20 December 1998 / Revised version: 7 May 1999 / Accepted: 27 May 1999     

Some like it cold: summer torpor by freetail bats in the Australian arid zone

Bats are among the most successful groups of Australian arid-zone mammals and, therefore, must cope with pronounced seasonal fluctuations in ambient temperature (T _a), food availability and unpredictable weather patterns. As knowledge about the energy conserving strategies in desert bats is scant, we used temperature-telemetry to quantify the thermal physiology of tree-roosting inland freetail bats (Mormopterus species 3, 8.5 g, n = 8) at Sturt National Park over two summers (2010–2012), when T _a was high and insects were relatively abundant. Torpor use and activity were affected by T _a. Bats remained normothermic on the warmest days; they employed one “morning” torpor bout on most days and typically exhibited two torpor bouts on the coolest days. Overall, animals employed torpor on 67.9 % of bat-days and torpor bout duration ranged from 0.5 to 39.3 h. At any given T _a, torpor bouts were longer in Mormopterus than in bats from temperate and subtropical habitats. Furthermore, unlike bats from other climatic regions that used only partial passive rewarming, Mormopterus aroused from torpor using either almost entirely passive (68.9 % of all arousals) or active rewarming (31.1 %). We provide the first quantitative data on torpor in a free-ranging arid-zone molossid during summer. They demonstrate that this desert bat uses torpor extensively in summer and often rewarms passively from torpor to maximise energy and water conservation.     

Haemolymph oxygen transport and acid-base status in Glyptonotus antarcticus Eights

Haemolymph samples were withdrawn from routinely active male intermoult Glyptonotus held at 0 ± 0.5°C, and analysed for blood-gas and acid-base variables. In both the arterialised (a) and venous (v) haemolymph, over 50% of the oxygen was transported as dissolved oxygen at PaO₂ and PvO₂ levels of 12.0 ± 1.15 and 7.70 ± 1.89 kPa, respectively. The maximum oxygen-carrying capacity of the haemocyanin (C^maxHcO₂) was relatively low at 0.19 ± 0.05 mmol l⁻¹, accompanied by relatively low protein and [Cu²⁺] levels indicating low circulating haemocyanin concentrations. Arterialised haemolymph had a mean pH of 7.88 ± 0.02(6) at a PCO₂ of 0.12 ± 0.01(6) kPa and a bicarbonate level of 12.95 ± 0.80(6) mequiv l⁻¹ with small differences in PCO₂ and pH between arterial and venous haemolymph. The non-bicarbonate buffering capacity of Glyptonotus haemolymph was low at −2.0 mequiv l⁻¹ HCO₃ ⁻ pH unit⁻¹. Haemolymph [l-lactate] and [d-glucose] levels were similar at < 1 mmol l⁻¹ in animals held in the laboratory and those sampled in Antarctica. The blood-gas and acid-base status of Glyptonotus haemolymph may be a reflection of the low and stable temperatures experienced by this Antarctic crustacean. Received: 14 August 1996 / Accepted: 3 November 1996     

Internal and External Influences on Near-Surface Microbial Community Structure in the Vicinity of the Cape Verde Islands

Microbial community structure in the subtropical north-east Atlantic Ocean was compared between 2 years and variation attributed to environmental variables. Surface seawater communities were analysed by flow cytometry and fluorescence in situ hybridisation. Probes specific to Alphaproteobacteria, Cyanobacteria, Gammaproteobacteria and Bacteroidetes identified 67–100% of cells. Due to natural variation in the study region due to the occurrence of major currents and islands, data could not be pooled but were instead divided between distinct water masses. Community structure did not differ greatly around the Cape Verde Islands between sampling periods but varied substantially in the open ocean, suggesting different environmental perturbations favour specific bacterial groups. Wind speed varied significantly between years, with moderate to strong breeze in winter 2008 and gales in winter 2006 (8.9 ± 0.2 ms⁻¹ and 16.0 ± 0.4 ms⁻¹, respectively). Enhanced wind-driven turbulence was associated with domination by the SAR11 clade of Alphaproteobacteria, which were present at 2.4-fold in the abundance of Prochlorococcus (41.8 ± 1.6% cells, compared to 17.7 ± 7.1%). Conversely, the calmer conditions of 2008 seemed to favour Prochlorococcus (40.0 ± 1.2% cells). Prochlorococcus high-light adapted clade HLI were only numerous during wind-driven turbulence, whereas oligotrophic-adapted clade HLII dominated under calm conditions. Bacteroidetes were most prominent in turbulent conditions (9.5 ± 1.3% cells as opposed to 4.7 ± 0.3%), as were Synechococcus. In 2008, a considerable dust deposition event occurred in the region, which may have led to the substantial Gammaproteobacteria population (22.5 ± 4.0% cells compared to 4.6 ± 0.6% in 2006). Wind-driven turbulence may have a significant impact on microbial community structure in the surface ocean. Therefore, community change following dust storm events may be linked to associated wind in addition to dust-derived nutrients.     

Phytoplankton biomass and production during late austral spring (1991) and summer (1993) in the Bransfield Strait

Phytoplankton biomass and productivity were measured during two cruises in the Bransfield Strait in December 1991 (D91) and January/February 1993 (J93). Strong seasonal variability in productivity values was observed due to differences in the physiological response of phytoplankton. However, although the photosynthetic capacity of phytoplankton was markedly lower in D91 [P _m ^B =0.61 ± 0.25 mg C (mg Chla)⁻¹ h⁻¹] than in J93 [P _m ^B =2.18 ± 0.91 mg C (mg Chla)⁻¹ h⁻¹], average water column chlorophyll values in different areas of the strait were approximately similar in D91 (49–78 mg Chla m⁻²) and J93 (22–76 mg Chla m⁻²). The spatial distribution of chlorophyll was patchy and generally associated with the influence of the different water masses that meet together in the Bransfield Strait. No correlation was found between the mixed layer depth and either the integrated chlorophyll or the productivity. Our results suggest that major phytoplankton blooms in the Bransfield Strait are advected from the nearby Gerlache Strait or Bellingshausen Sea following the main eastward surface currents. Accepted: 5 July 1998