Insulation and thermal balance of fasting harp and grey seal pups.

1. Harp and grey seal pups were examined during the post-weaning period to quantify their thermoregulatory abilities and thermal limits. 2. Deep body temperatures of harp seals (37.8 +/- 0.8 degrees C) were not significantly different from those of grey seals (38.9 +/- 0.4 degrees C). 3. As blubber depth declined during the fast, temperature gradients extended increasingly deeper into the muscle layer potentially decreasing heat loss. 4. Blubber conductivity (approximately 0.18 W/m/degrees C) did not vary regionally within an animal, or between animals or species. 5. Calculated lethal cold limits in air were between -85.4 degrees C and -116.1 degrees C, suggesting that fasting, weaned pups can easily cope with temperatures they would normally experience.     

MEASURING CHANGES IN JUVENILE GRAY SEAL BODY COMPOSITION

Determining body composition in gray seals ( Halichoerus grypus ) is important when studying their physiology and life history. In this study we investigated the predictability of total body fat (TBF) and protein (TBP) in postweaned gray seal pups from morphometric measurements, blubber thickness using ultrasonographs and bioelectrical impedance analysis (BIA). In postweaned pups, TBF (kg) could be estimated from girth measurements and sex ( n = 45, r ²= 0.878) using hydrogen isotope dilution methods as the reference. However, TBP could not be reliably estimated from morphometric data. TBF (kg) in yearlings was best predicted from mass ( n = 6, r ²= 0.776) and TBP (kg) from mass/length ( r ²= 0.949). Dorsal blubber thickness using B-mode ultrasound was also a significant predictor of TBF (kg) in postweaned pups ( r ²= 0.725) but BIA was not. Marked pups were recaptured during their first few months of life ( n = 48) and body composition changes investigated. Animals lost mass and TBF after leaving the breeding beach, largely during the first 5–6 mo of life. Postweaned pups were ∼40% TBF and ∼13% TBP whereas yearlings were ∼12% TBF and ∼20% TBP. Pups that survived beyond 6 mo of age then regained mass as protein. Morphometric measurements are a useful field indication of body condition when isotope dilution is impractical.     

Observations on the infectivity of parasitic third-stage larvae of Uncinaria lucasi Stiles 1901 (Nematoda: Ancylostomatidae) of Northern fur seals, Callorhinus ursinus Linn., on St. Paul Island, Alaska.

Twelve fur seal pups, which had not nursed their mothers, were used in an infectivity experiment. Pups were exposed to parasitic 3rd-stage larvae of Uncinaria lucasi from belly tissues of fur seal bulls, bachelors, and pregnant cows, to determine maturation capability of the larvae. Hookworms were not recovered from the intestines of 3 pups receiving larvae from belly blubber of bulls, 6 pups receiving larvae from belly blubber of bachelors, and 1 nonexposed pup. Maturation of hookworms did occur in 2 pups exposed to larvae from a mixture of belly blubber, mammary tissue, and milk of pregnant cows. Parasitic 3rd-stage hookworm larvae from belly tissues of pregnant and "non-pregnant" fur seal cows averaged 938.1 and 802.1 micron long, and 34.1 and 31.5 micron wide, respectively; however, larvae from belly tissues of a fur seal bull, bachelors, 2-year-old males, male and female yearlings and pups, and Steller Sea Lion subadults averaged 640.5-732.0 micron long and 20.9-24.9 micron wide.     

Ecological implications of body composition and thermal capabilities in young antarctic fur seals (Arctocephalus gazella)

In comparison with other homeotherms, young recently weaned marine mammals in high latitudes face exceptional energetic demands when foraging and thermoregulating. Lipids are an important source of energy and a major component of insulation that allows them to meet these demands. To examine the role of lipid stores in a high-latitude pinniped, we measured the body composition and thermoregulatory capabilities of Antarctic fur seal (Arctocephalus gazella) pups and yearlings by using flow-through respirometry and hydrogen isotope dilution. From these data, we constructed a model to examine the importance of postweaning fasting capability in free-ranging young fur seals. Resting metabolic rates were different for pups and yearlings measured in 0.6 degrees C water, 10.3 degrees C water, and ambient air; however, mass and percent lipid as covariates accounted for the different metabolic responses in pups and yearlings for all treatments. The estimated lower critical temperature for combined pups and yearlings was 14.4 degrees C, 10 degrees -15 degrees C above water temperatures normally experienced by Antarctic fur seals. Modeling predicted that a weaned fur seal pup would survive at sea from 9.8 to 36.2 d before succumbing to starvation. The most likely maximum travel distance within this time constraint suggests that food resources close to the natal rookery are important to first-year survival for this species.     

Development of body oxygen stores in harbor seals: effects of age, mass, and body composition

Harbor seal pups are highly precocial and can swim and dive at birth. Such behavioral maturity suggests that they may be born with mature body oxygen stores or that stores develop quickly during the nursing period. To test this hypothesis, we compared the blood and muscle oxygen stores of harbor seal pups, yearlings, and adults. We found that pups had smaller oxygen stores than adults (neonates 57%, weaned pups 75%, and yearlings 90% those of adults), largely because neonatal myoglobin concentrations were low (1.6+/-0.2 g% vs. 3.8+/-0.3 g% for adults) and changed little during the nursing period. In contrast, blood oxygen stores were relatively mature, with nursing pups having hematocrit (55%+/-0.2%), hemoglobin (21.7+/-0.4 g%), and blood volume (12.3+/-0.5 mL/kg) only slightly lower than the corresponding values for adults (57%+/-0.2%, 23.8+/-0.3 g %, and 15.0+/-0.5 mL/kg). Because neonatal pups had relatively high metabolic rates (11.0 mL O2/kg min), their calculated aerobic dive limit was less than 50% that of adults. These results suggest that harbor seals' early aquatic activity is primarily supported by rapid development of blood, with immature muscle oxygen stores and elevated use rates limiting aerobic diving ability.     

Harbor seal pup dispersal and individual morphology,hematology, and contaminant factors affecting survival

Understanding the factors affecting individual harbor seal (Phoca vitulina) survival is essential for determining population level health risks. We estimated postweaning dispersal, and modeled the effects of morphology, hematology, and blubber contaminants on the survival of recently weaned harbor seal pups using a mark recapture framework. We deployed satellite transmitters on apparently healthy pups captured in San Francisco Bay (SFB, n = 19) and Tomales Bay (TB, n = 7), and pups released after rehabilitation that stranded along the central California coast preweaning (n = 21). Dispersal distances were further than previously reported for harbor seal pups (maximum = 802 km) which has implications for understanding risks to this vulnerable age class. We found differences in body condition, serum immunoglobulin and thyroxine (T4) concentrations, white blood cell count, and blubber organohalogen contamination (OH) among the three groups. Overall, increased T4, decreased OH, and increased mass were associated with greater survival probabilities; whereas, among stranded seals, greater mass gain, shorter time in rehabilitation, and admission to rehabilitation earlier in the season were associated with greater survival probabilities. Attention to these latter factors may improve the success of rehabilitation efforts. For wild pups, reduction of legacy contaminants and direct causes of mortality, such as ship strike, may enhance pup survival.     

A three-compartment thermometry model for the improved estimation of changes in body heat content

The aim of this study was to use whole body calorimetry to directly measure the change in body heat content (DeltaH(b)) during steady-state exercise and compare these values with those estimated using thermometry. The thermometry models tested were the traditional two-compartment model of "core" and "shell" temperatures, and a three-compartment model of "core," "muscle," and "shell" temperatures; with individual compartments within each model weighted for their relative influence upon DeltaH(b) by coefficients subject to a nonnegative and a sum-to-one constraint. Fifty-two participants performed 90 min of moderate-intensity exercise (40% of Vo(2 peak)) on a cycle ergometer in the Snellen air calorimeter, at regulated air temperatures of 24 degrees C or 30 degrees C and a relative humidity of either 30% or 60%. The "core" compartment was represented by temperatures measured in the esophagus (T(es)), rectum (T(re)), and aural canal (T(au)), while the "muscle" compartment was represented by regional muscle temperature measured in the vastus lateralis (T(vl)), triceps brachii (T(tb)), and upper trapezius (T(ut)). The "shell" compartment was represented by the weighted mean of 12 skin temperatures (T(sk)). The whole body calorimetry data were used to derive optimally fitting two- and three-compartment thermometry models. The traditional two-compartment model was found to be statistically biased, systematically underestimating DeltaH(b) by 15.5% (SD 31.3) at 24 degrees C and by 35.5% (SD 21.9) at 30 degrees C. The three-compartment model showed no such bias, yielding a more precise estimate of DeltaH(b) as evidenced by a mean estimation error of 1.1% (SD 29.5) at 24 degrees C and 5.4% (SD 30.0) at 30 degrees C with an adjusted R(2) of 0.48 and 0.51, respectively. It is concluded that a major source of error in the estimation of DeltaH(b) using the traditional two-compartment thermometry model is the lack of an expression independently representing the heat storage in muscle during exercise.     

DIET OF WEANED PUPS AND SEASONAL VARIATIONS IN BODY CONDITION OF JUVENILE BARENTS SEA HARP SEALS PHOCA GROENLANDICA

Data on blubber depth, body condition, and diet were collected from young of year and juvenile harp seals ( Phoca groenlandica ) in the Barents Sea in the periods February-April and June-October 1990–1997. Harp seal pups feed independently shortly after weaning. In the southern parts of the Barents Sea the diet of harp seal pups consisted mainly of euphausiids ("krill") Thysanoessa sp. and amphipods of the genus Parathemisto . The change in body condition of harp seal pups between weaning (mid-March) and June indicated that considerable amounts of blubber energy, deposited during suckling, were mobilized. This suggests that the early food intake of weaned pups was insufficient to meet the energy requirements. The pattern of low spring and earlysummer body condition, followed by a subsequent rapid improvement in condition during late summer and autumn appeared to occur not only among pups-of-the-year, but also in one- and two-year old harp seals.     

Thermoregulatory responses to cold water at different times of day.

This study examined how time of day affects thermoregulation during cold-water immersion (CWI). It was hypothesized that the shivering and vasoconstrictor responses to CWI would differ at 0700 vs. 1500 because of lower initial core temperatures (T(core)) at 0700. Nine men were immersed (20 degrees C, 2 h) at 0700 and 1500 on 2 days. No differences (P > 0.05) between times were observed for metabolic heat production (M, 150 W. m(-2)), heat flow (250 W. m(-2)), mean skin temperature (T(sk), 21 degrees C), and the mean body temperature-change in M (DeltaM) relationship. Rectal temperature (T(re)) was higher (P < 0.05) before (Delta = 0.4 degrees C) and throughout CWI during 1500. The change in T(re) was greater (P < 0. 05) at 1500 (-1.4 degrees C) vs. 0700 (-1.2 degrees C), likely because of the higher T(re)-T(sk) gradient (0.3 degrees C) at 1500. These data indicate that shivering and vasoconstriction are not affected by time of day. These observations raise the possibility that CWI may increase the risk of hypothermia in the early morning because of a lower initial T(core).     

Effect of high-load and high-volume resistance exercise on the tensiomyographic twitch response of biceps brachii

The purpose of the present study was to assess the ability of TMG in detecting mechanical fatigue induced by two different resistance exercises on biceps brachii: high-volume (HV), and high-load (HL). Sixteen healthy subjects (age 25.1±2.6years; body mass 79.9±8.9kg; height 179±7.4cm) performed arm-curl in two different protocols (HV: 8×15×10kg, HL: 5×3×30kg). Tensiomyography was used to assess muscle response to both exercise protocols. The contractile capacity of biceps brachii significantly varied by means of the effects of potentiation and fatigue mechanisms that take place at different exercise phases. The most significant changes correspond to values of maximum radial displacement of muscle belly (D(m)), sustained contraction time (T(s)), relaxation time (T(r)), and contraction velocity (V(c)). The behavior of these parameters is, in general, similar in both exercise protocols, but they show subtle differences among them. During the first set, in both protocols, values for V(c) increase, along with a decrease in T(r), T(s), and D(m) values. Fatigue onset was evident from changes in such parameters, with HL being the first in showing these mechanisms. Tensiomyography has been shown to be highly sensitive in detecting fatigue-induced changes.     

BODY TEMPERATURE IN THE NEWBORN SOUTHERN ELEPHANT SEAL, MIROUNGA LEONINA, AT MACQUARIE ISLAND

Newborn southern elephant seal pups were reported by Laws (1953) to be "to some extent poikilothermic at birth." Rectal temperatures of known age southern elephant seal pups were recorded during the 1985 pupping season at Macquarie Island. The mean pup rectal temperature was found to be 381°C ± 0.1°C SEM ( n = 131, range = 36.5°-39.1°C). Pups at two hours, six hours, and one day after birth had significantly higher rectal temperatures than pups two, three, or four days of age. Rectal temperatures of neonatal southern elephant seals were within the range observed for other pinnipeds, (but never as low as the 31°C previously observed for southern elephant seals at Signy Island in 1953). A significant though weak positive correlation was found between pup temperature and body weight. However, no correlation between pup temperature and age or any environmental factor was found. These observations demonstrated that southern elephant seal pups at Macquarie Island are homeothermic, rather than heterothermic from birth.     

Shifts in thermoregulatory strategy during ontogeny in harp seals (Pagophilus groenlandicus)

Heat balance can be difficult for young and/or small animals in polar regions because environmental conditions in combination with small body size or physiological immaturity can increase heat loss. We investigated how thermoregulatory patterns change with ontogeny in 5 age classes of harp seal (Pagophilus groenlandicus) from birth to post-molt to further understand the timing of thermoregulatory development in relation to their potential vulnerability to ongoing fluctuations in the extent and stability of Arctic pack ice. We measured changes in the amount, conductivity, and resistance of the seal pups׳ insulative layers (blubber and fur), the potential for endogenous heat-generation by shivering (muscle enzyme activity), and nonshivering thermogenesis (NST; brown adipose tissue (BAT) uncoupling protein 1 (UCP1) expression and mitochondrial density). There was no significant difference in blubber conductivity among age classes, though the amount of blubber insulation significantly increased from birth to weaning. Pelage conductivity was low (0.12±0.01 W m⁻¹ °C⁻¹) except in 9-day old pups (0.40±0.08 W m⁻¹ °C⁻¹); the significantly higher conductivity may signal the beginning of the molt, and this age group may be the most vulnerable to early water entry. Citrate synthase activity significantly increased (49.68±3.26 to 75.08±3.52 μmol min⁻¹ g wet weight⁻¹) in the muscle; however it is unlikely that increasing a single enzyme greatly impacts heat generation. BAT of younger pups contained UCP1, though expression and mitochondrial density quickly declined, and the ability of pups to produce heat via NST was lost by weaning. While total thermal resistance did not differ, neonatal and early nursing animals gained the majority of their thermal resistance from lanugo (82.5±0.03%); however, lanugo is not insulative when wet, and NST may be important to maintain euthermia and dry the coat if early immersion in water occurs. By late nursing, blubber seems sufficient as insulation (75.87±0.01% of resistance after 4 weeks), but high conductivity of fur may be responsible for retention of UCP1 expression. Weaned animals rely on blubber insulation, and no longer need NST, as wetted fur is no longer a threat to euthermia.     

The effect of prey quality and ice conditions on the nutritional status of Baltic gray seals of different age groups

We analyzed a long-term data set of the body condition of Baltic gray seals (Halichoerus grypus) over time and investigated how average subcutaneous blubber thickness of different age groups of seals corresponds to environmental factors. Blubber thickness of pups declined until 2010. The decreasing weight of 5–6-year-old herring (Clupea harengus), the main prey fish for Baltic gray seals, explained well the decline. In the Gulf of Finland, the blubber thickness of pups declined also in recent years (2011–2015) with declining number of days with permanent ice cover. In other regions, the blubber thickness of pups increased during recent years with increasing weight of herring. The blubber thickness of sub-adults in Baltic Proper and that of hunted adult females in the Bothnian Bay also increased during recent years, and the weight of age 6+ or 7-year-old herring best explained the increase. The blubber thickness of all age groups of seals was thinnest in the Bothnian Bay where also herring weight was lowest. There was a negative correlation between blubber thickness of seals and herring catch size (an index of herring abundance) suggesting that herring quality, not the quantity, is important for the nutritional status of Baltic gray seals. Nutritional status of gray seals may thus reveal changes in the marine food web which affect herring quality. Marine food web, in turn, may be affected, e.g., by climate change. The warming climate also has an impact on ice cover and thus body condition of seal pups.     

Sex-based differences and similarities in locomotor performance, thermal preferences, and escape behaviour in the lizard Platysaurus intermedius wilhelmi

Differences between sexes in physiological performance have received little attention in animals. We tested for sex differences in maximum sprint speed and maximal exertion over a range of temperatures in a population of Platysaurus intermedius wilhelmi lizards. We also examined sex-based differences in selected temperature range, mean field body temperatures (T(b)), and thermal activity limits. Finally, we conducted field studies to quantify male and female responses to a potential predator, which may be affected by their respective performance capabilities. Males were faster than females at all temperatures, and body size had no significant effect on sprint speeds. Males and females also selected similar T(b)'s when placed in a thermal gradient, but in the field, male lizards' T(b)'s were different from those of the females. However, predicted sprint speeds for males and females at their field T(b)'s are similar. No significant differences were found between males and females with regard to maximal exertion. When approached in the field, adult male lizards took refuge significantly earlier than did adult females and also fled over shorter distances, suggesting that females rely on crypsis as an escape strategy.     

THE RELIABILITY OF SKINFOLD-CALIPERS FOR MEASURING BLUBBER THICKNESS OF STELLER SEA LION PUPS (EUMETOPIAS JUBATUS)

Twelve dead Steller sea lion pups ( Eumetopias jubatus ) aged 3-14 d were recovered from rookeries in Southeast Alaska. They had a wide range of body sizes and conditions (small to large and fat to no fat). The ability of calipers to estimate the thickness of their blubber layer was assessed with a set of skinfold calipers. Average error of measurement for skin and blubber thickness was an acceptable 5.4%, but the skin and blubber of the pups were highly compressible. Skinfold thickness increased with body mass but did not necessarily reflect the development of blubber, given that pups with no blubber also showed an increase in skinfold thickness with increases in body mass. Skinfold thickness of sea lion pups appears to predict body size better than it predicts blubber thickness, making it difficult if not impossible to develop a simple index of body condition or a calculation of percent body fat for Steller sea lion pups from skinfold caliper measurements.     

THERMAL FUNCTION OF PHOCID SEAL FUR

In phocid seals, blubber serves as the main thermal insulation instead of fur. The thermal function of fur, at least in adult phocid seals, has therefore been questioned. We measured the relative contribution of fur to the combined thermal resistance (insulation) offered by blubber, skin, and fur in newborn and adult harp ( Pagophilus groenlandicus ) and hooded ( Cystophora cristata ) seals, in air and water, to elucidate the role of fur as insulation in phocid seals. In air the fur contributed 90% of the combined thermal resistance of blubber, skin, and fur in newborn harp seal pups and 29% in adulrs, whereas in hooded seals the fur contributed 73% in newborn pups and 34% in adults. When submerged the thermal resistance of the fur was reduced by 84%-92%, and contributed 65% to the total insulation in newborn harp seal pups and 3% in adults, and 26% in newborn hooded seal pups and 5% in adults. We conclude that in air the fur of phocid seals makes an important contribution to the insulation of pups, and also contributes considerably to the insulation of adult animals. In water, even though the thermal resistance of the fur is dramatically reduced, the fur still contributes substantially to the insulation of pups, but its contribution in adults is negligible.     

Thermoregulation of weaned northern elephant seal (Mirounga angustirostris) pups in air and water

Fasting weaned northern elephant seal pups (Mirounga angustirostris) experience diverse environmental conditions on land and in water on a daily basis. Each environment undoubtedly induces distinct energetic costs that may vary for pups of differing body condition. To determine the energetic costs associated with different environmental conditions and whether costs vary between individuals, body mass, surface area, volume, body composition, resting metabolic rate, and core body temperature were determined for 17 weaned northern elephant seal pups from A?o Nuevo, California. Metabolic rate and body temperature were measured for pups resting in air (20.9 degrees +/-0.8 degrees C), cold water (3.8 degrees+/-0.4 degrees ;C), and warm water (14.5 degrees+/-0.2 degrees C). Resting metabolic rate increased with body mass (range: 62.0-108.0 kg) and was also correlated with lean mass and lipid mass. Metabolic rates ranged from 293.6 to 512.7 mL O(2) min(-1) and were lowest for pups resting in cold water. Thermal conductance, calculated from metabolic rate and core body temperature, ranged from 3.1 to 15.2 W degrees C(-1), with the highest values in air and the lowest values in cold water. Metabolic responses to the three environmental conditions did not differ with individual variation in body condition. For all elephant seal pups, a consequence of high lipid content is that thermoregulatory costs are greatest on land and lowest in cold water, a pattern that contrasts markedly with terrestrial mammals.     

Exercise in the heat is limited by a critical internal temperature.

We examined whether fatigue during exertional heat stress occurred at a critical internal temperature independent of the initial temperature at the start of exercise. Microwaves (2.1 GHz; 100 mW/cm(2)) were used to rapidly (3-8 min) heat rats before treadmill exercise to exhaustion. In a repeated-measures design, food-restricted male Sprague-Dawley rats (n = 11) were preheated to three levels (low, medium, and high). In addition, two sham exposures, Sham 1 and Sham 2, were administered at the beginning and end of the study, respectively. At the initiation of exercise, hypothalamic (T(hyp)) and rectal (T(rec)) temperatures ranged from 39.0 degrees C to 42.8 degrees C (T(hyp)) and 42.1 degrees C (T(rec)). The treadmill speed was 17 m/min (8 degrees grade), and the ambient temperature during exercise was 35 degrees C. Each treatment was separated by 3 wk. Run time to exhaustion was significantly reduced after preheating. There was a significant negative correlation between run time and initial T(hyp) and T(rec) (r = 0.73 and 0.74, respectively). The temperatures at exhaustion were not significantly different across treatments, with a range of 41.9-42.2 degrees C (T(hyp)) and 42.2-42.5 degrees C (T(rec)). There were no significant differences in run time in the sham runs administered at the start and end of the investigation. No rats died as a result of exposure to any of the treatments, and body weight the day after each treatment was unaffected. These results support the concept that a critical temperature exists that limits exercise in the heat.     

SEASONAL CHANGES IN BLUBBER DISTRIBUTION IN ATLANTIC HARBOR SEALS: INDICATIONS OF THERMODYNAMIC CONSIDERATIONS

Among its functions, the hypodermal blubber layer of pinnipeds serves as both an energy reserve and insulation. This study examined seasonal changes in blubber distribution and body morphology in a group of captive changesharbor seals to test whether these changes were designed to maximize insulative effectiveness. Seasonal changes were found in girth, blubber volume, mean blubber depth, and the ratio of blubber depth to body radius (d/r ratio). These changes were more evident in older seals. The d/r ratio demonstrated a smaller relative annual change than mean blubber depth. The d/r ratio also exhibited less variation along the length of the seal than blubber depth at any given time. Similar to reports for ringed seals, and contrary to those for southern elephant seals, the harbor seals preferentially lost blubber from overinsulated areas of the body. These results suggest that core tissue and blubber mass are lost in a manner that maximizes insulative effectiveness.     

Body temperature regulation in the rat

In loosely-restrained adult conscious rats exposed to stepwise changes in ambient temperature (T(a)) from 25 to 5 degrees C or from 20 to 35 degrees C, we have recorded body and tail temperatures, metabolic rate (VO(2)), shivering and ventilation (V). It was found that VO(2) and V vary with T(a) and show a nadir for a T(a) of 30 degrees C whereas shivering starts at 20 degrees C and increases progressively with cold exposure. T(tail) follows changes in T(a) whereas T(body) decreases slightly in cold and increases markedly in warm exposure. These results suggest that the control of T(body) interacts with the control of breathing in order to increase VO(2) during cold exposure and to facilitate evaporative respiratory heat dissipation during warm exposure.