Tufted ducks Aythya fuligula do not control buoyancy during diving

Work against buoyancy during submergence is a large component of the energy costs for shallow diving ducks. For penguins, buoyancy is less of a problem, however they still seem to trade‐off levels of oxygen stores against the costs and benefits of buoyant force during descent and ascent. This trade‐off is presumably achieved by increasing air sac volume and hence pre‐dive buoyancy (B_pre) when diving deeper. Tufted ducks, Aythya fuligula, almost always dive with nearly full oxygen stores so these cannot be increased. However, the high natural buoyancy of tufted ducks guarantees a passive ascent, so they might be expected to decrease B_pre before particularly deep, long dives to reduce the energy costs of diving. Body heat lost to the water can also be a cause of substantial energy expenditure during a dive, both through dissipation to the ambient environment and through the heating of ingested food and water. Thus dive depth (d_d), duration and food type can influence how much heat energy is lost during a dive. The present study investigated the relationship between certain physiological and behavioural adjustments by tufted ducks to d_d and food type. Changes in B_pre, deep body temperature (T_b) and dive time budgeting of four ducks were measured when diving to two different depths (1.5 and 5.7 m), and for two types of food (mussels and mealworms). The hypothesis was that in tufted ducks, B_pre decreases as d_d increases. The ducks did not change B_pre in response to different diving depths, and thus the hypothesis was rejected. T_b was largely unaffected by dives to either depth. However, diving behaviour changed at the greater d_d, including an increase in dive duration and vertical descent speed. Behaviour also changed depending on the food type, including an increase in foraging duration and vertical descent speed when mussels were present. Behavioural changes seem to represent the major adjustment made by tufted ducks in response to changes in their diving environment.     

The influence of oxygen and carbon dioxide on diving behaviour of tufted ducks, Aythya fuligula

While optimal diving models focus on the diver's oxygen (O(2)) stores as the predominant factor influencing diving behaviour, many vertebrate species surface from a dive before these stores are exhausted and may commence another dive well after their O(2) stores have been resaturated. This study investigates the influence of hypoxia and also hypercapnia on the dive cycle of tufted ducks, Aythya fuligula, in terms of surface duration and dive duration. The birds were trained to surface into a respirometer box after each dive to a feeding tray so that rates of O(2) uptake (VO2) and carbon dioxide output (VCO2) at the surface could be measured. Although Vco2 initially lagged behind Vo2, both respiratory gas stores were close to full adjustment after the average surface duration, indicating that they probably had a similar degree of influence on surface duration. Chemoreceptors, which are known to influence diving behaviour, detect changes in O(2) and CO(2) partial pressures in the arterial blood. Thus, the need to restore blood gas levels appears to be a strong stimulus to continue ventilation. Mean surface duration coincided with peak instantaneous respiratory exchange ratio due to predive anticipatory hyperventilation causing hypocapnia. For comparison, the relationship between surface duration and O(2) uptake in reanalysed data for two grey seals indicated that one animal tended to dive well after fully restocking its O(2) stores, while the other dived at the point of full restocking. More CO(2) is exchanged than O(2) in tufted ducks during the last few breaths before the first dive of a bout, serving to reduce CO(2) stores and suggesting that hypercapnia rather than hypoxia is more often the limiting factor on asphyxia tolerance during dives. Indeed, according to calculations of O(2) stores and O(2) consumption rates over modal diving durations, a lack of O(2) does not seem to be associated with the termination of a dive in tufted ducks. However, factors other than CO(2) are also likely to be important, and perhaps more so, such as food density and rate of food ingestion. Because some predictive success has been demonstrated for optimal diving models, they should continue to incorporate O(2) stores as a variable, but their validity is likely to be improved by also focusing on CO(2) stores.     

Energetic costs of surface swimming and diving of birds

The energetic costs of swimming at the surface (swimming) and swimming underwater (diving) are compared in tufted ducks (Aythya fuligula) and three species of penguins, the gentoo (Pygoscelis papua), the king (Aptenodytes patagonicus), and the emperor (Aythya forsteri). Ducks swim on the surface and use their webbed feet as paddles, whereas penguins tend to swim just below the surface and use their flippers as hydrofoils, the latter being much more efficient. Penguins are more streamlined in shape. Thus, the amount of energy required to transport a given mass of bird a given distance (known as the cost of transport) is some two to three times greater in ducks than in penguins. Ducks are also very buoyant, and overcoming the force of buoyancy accounts for 60% and 85% of the cost of descent and remaining on the bottom, respectively, in these birds. The energy cost of a tufted duck diving to about 1.7 m is similar to that when it is swimming at its maximum sustainable speed at the surface (i.e., approximately 3.5 times the value when resting on water). Nonetheless, because of the relatively short duration of its dives, the tufted duck dives well within its calculated aerobic dive limit (cADL, usable O(2) stores per rate of O(2) usage when underwater). However, these three species of penguins have maximum dive durations ranging from 5 min to almost 16 min and maximum dive depths from 155 to 530 m. When these birds dive, they have to metabolise at no more than when resting in water in order for cADL to encompass the duration of most of their natural dives. In gentoo and king penguins, there is a fall in abdominal temperature during bouts of diving; this may reduce the oxygen requirements in the abdominal region, thus enabling dive duration to be extended further than would otherwise be the case.     

To what extent is the foraging behaviour of aquatic birds constrained by their physiology?

Aquatic birds have access to limited amounts of usable oxygen when they forage (dive) underwater, so the major physiological constraint to their behaviour is the need to periodically visit the water surface to replenish these stores and remove accumulated carbon dioxide. The size of the oxygen stores and the rate at which they are used (V dot o2) or carbon dioxide accumulates are the ultimate determinants of the duration that aquatic birds can remain feeding underwater. However, the assumption that the decision to terminate a dive is governed solely by the level of the respiratory stores is not always valid. Quantification of an optimal diving model for tufted ducks (Aythya fuligula) shows that while they dive efficiently by spending a minimum amount of time on the surface to replenish the oxygen used during a dive, they dive with nearly full oxygen stores and surface well before these stores are exhausted. The rates of carbon dioxide production during dives and removal during surface intervals are likely to be at least as important a constraint as oxygen; thus, further developments of optimal diving models should account for their effects. In the field, diving birds will adapt to changing environmental conditions and often maximise the time spent submerged during diving bouts. However, other factors influence the diving depths and durations of aquatic birds, and in some circumstances they are unable to forage sufficiently well to provide food for their offspring. The latest developments in telemetry have demonstrated how diving birds can make physiological decisions based on complex environmental factors. Diving penguins can control their inhaled air volume to match the expected depth, likely prey encounter rate, and buoyancy challenges of the following dive.     

Implications of river damming: the influence of aquatic hypoxia on the diving physiology and behaviour of the endangered Mary River turtle

River impoundments are characterized by low oxygen levels as a result of reduced water velocity and increased water depth. Bimodally respiring turtle species are likely to be highly sensitive to changes in aquatic PO₂ with decreases in oxygen levels impacting upon their diving ability. The acute and long-term effects of aquatic hypoxia on dive duration, oxygen consumption and blood respiratory properties were examined in hatchlings of the endangered Mary River turtle Elusor macrurus . It was hypothesized that acute exposure to aquatic hypoxia would cause a decrease in dive duration as a consequence of a decrease in reliance on aquatic respiration. With long-term exposure to hypoxia, we predicted that Elu. macrurus would have the capacity to compensate for the acute effect of hypoxia and that dive duration would increase due to an increase in aquatic respiration, haemoglobin concentration and oxygen affinity (P₅₀). When exposed to hypoxic conditions, aquatic respiration in Elu. macrurus was substantially reduced resulting in a 51% decrease in dive duration. Contrary to our predictions, Elu. macrurus hatchlings did not acclimate, and long-term exposure to hypoxic conditions caused Elu. macrurus to lose significantly more oxygen to the hypoxic water than the normoxic acclimated turtles. The exacerbation of long-term hypoxia on the respiratory physiology and diving ecology of Elu. macrurus raises concerns about the impacts of long-term environmental change as a result of habitat alteration on the survival of freshwater turtle populations.     

Testing optimal foraging models for air-breathing divers

Models of diving optimality qualitatively predict diving behaviours of aquatic birds and mammals. However, none of them has been empirically tested. We examined the quantitative predictions of optimal diving models by combining cumulative oxygen uptake curves with estimates of power costs during the dives of six tufted ducks, Aythya fuligula. The effects of differing foraging costs on dive duration and rate of oxygen uptake (V_O2up) at the surface were measured during bouts of voluntary dives to a food tray. The birds were trained to surface into a respirometer after each dive, so that changes in V_O2up over time could be measured. The tray held either just food or closely packed stones on top of the food to make foraging energetically more costly. In contrast to predictions from the Houston & Carbone model, foraging time (t_f) increased after dives incorporating higher foraging energy costs but surface time (t_s) remained the same. While optimal diving models have assumed that the cumulative oxygen uptake curve is fixed, V_O2up increased when the energy cost of the dive increased. The optimal breathing model quantitatively predicted ts in both conditions and oxygen consumption during foraging (m₂t_f) in the control condition, for the mean of all ducks. This offers evidence that the ducks were diving optimally and supports the fundamentals of optimal diving theory. However, the model did not consistently predictt_s or m₂t_f for individual birds. We discuss the limits of optimal foraging models for air-breathing divers caused by individual variation. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

SUMMER DIVING BEHAVIOR OF MALE WALRUSES IN BRISTOL BAY, ALASKA

Pacific walruses ( Odobenus rosmarus divergens ) make trips from ice or land haul-out sites to forage for benthic prey. We describe dive and trip characteristics from time-depth-recorder data collected over a one-month period during summer from four male Pacific walruses in Bristol Bay, Alaska. Dives were classified into four types. Shallow (4 m), short (2.7 min), square-shaped dives accounted for 11% of trip time, and many were probably associated with traveling. Shallow (2 m) and very short (0.5 min) dives composed only 1% of trip time. Deep (41 m), long (7.2 min), square-shaped dives accounted for 46% of trip time and were undoubtedly associated with benthic foraging. V-shaped dives ranged widely in depth, were of moderate duration (4.7 min), and composed 3% of trip time. These dives may have been associated with navigation or exploration of the seafloor for potential prey habitat. Surface intervals between dives were similar among dive types, and generally lasted 1–2 min. Total foraging time was strongly correlated with trip duration and there was no apparent diel pattern of diving in any dive type among animals. We found no correlation between dive duration and postdive surface interval within dive types, suggesting that diving occurred within aerobic dive limits. Trip duration varied considerably within and among walruses (0.3–9.4 d), and there was evidence that some of the very short trips were unrelated to foraging. Overall, walruses were in the water for 76.6% of the time, of which 60.3% was spent diving.     

Diving pattern and performance in the macaroni penguin Eudptes chrysolophus

The pattern and characteristics of diving in two female macaroni penguins Eudyptes chrysolophus was studied, during the brooding period, using continuous-recording time-depth recorders, for a total of I8 days (15 consecutive days) during which the depth, duration and timing of 4876 dives were recorded. Diving in the first 11 days was exclusively diurnal, averaging 244 dives on trips lasting 12 hours. Near the end of the brooding period trips were longer and included diving at night. About half of all trips (except those involving continuous night-time diving) was spent in diving and dive rate averaged 14–25 dives per hour (42 per hour at night). The duration of day time dives varied between trips, and averaged 1.4–1.7 min, with a subsequent surface interval of 0.5–0.9 min. Dive duration was significantly directly related to depth, the latter accounting for 53% of the variation. The average depths of daytime dives were 20–35 m (maximum depth 11 5 m). Dives at night were shorter (average duration 0.9 min) and much shallower (maximum 11 m); depth accounted for only 6% of the variation in duration. Estimates of potential prey capture rates (3–5 krill per dive; one krill every 17–20 s) are made. Daily weight changes in chicks were directly related to number of dives, but not to foraging trip duration nor time spent diving. Of the other species at the same site which live by diving to catch krill, gentoo penguins forage exclusively diurnally, making longer. deeper dives; Antarctic fur seals, which dive to similar depths as macaroni penguins, do so mainly at night.     

The effects of breathing different levels of O2 and CO2 on the diving responses of ducks (Anas platyrhynchos) and cormorants (Phalacrocorax auritus)

Summary The effects of breathing different levels of O₂ and CO₂ before forced dives were investigated in 5 dabbling ducks (White Pekin) and 5 deep divers (Double Crested Cormorants). Breathing and heart rates, blood gases, and blood pH, were monitored. After breathing air before diving, ducks exhibited a slow decrease in heart rate that reached a minimum of 20 beats·min⁻¹ after 50 s submergence. The development of bradycardia was retarded if the duck breathed a hyperoxic gas mixture before diving and was accelerated if the gas mixture was hypoxic and hypercapnic. The cormorants' diving heart rate decreased to a minimum of about 60 beats·min⁻¹ in less than 20 s and development of bradycardia was unaffected by different levels of O₂ and CO₂ breathed before diving. Consequently, bradycardia in forced dived cormorants was unrelated to changes in blood gases in the dives which suggests that intravascular chemoreceptors are unimportant in initiating diving bradycardia in cormorants.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

The diving paradox: new insights into the role of the dive response in air-breathing vertebrates

When aquatic reptiles, birds and mammals submerge, they typically exhibit a dive response in which breathing ceases, heart rate slows, and blood flow to peripheral tissues is reduced. The profound dive response that occurs during forced submergence sequesters blood oxygen for the brain and heart while allowing peripheral tissues to become anaerobic, thus protecting the animal from immediate asphyxiation. However, the decrease in peripheral blood flow is in direct conflict with the exercise response necessary for supporting muscle metabolism during submerged swimming. In free diving animals, a dive response still occurs, but it is less intense than during forced submergence, and whole-body metabolism remains aerobic. If blood oxygen is not sequestered for brain and heart metabolism during normal diving, then what is the purpose of the dive response? Here, we show that its primary role may be to regulate the degree of hypoxia in skeletal muscle so that blood and muscle oxygen stores can be efficiently used. Paradoxically, the muscles of diving vertebrates must become hypoxic to maximize aerobic dive duration. At the same time, morphological and enzymatic adaptations enhance intracellular oxygen diffusion at low partial pressures of oxygen. Optimizing the use of blood and muscle oxygen stores allows aquatic, air-breathing vertebrates to exercise for prolonged periods while holding their breath.     

A review of the multi-level adaptations for maximizing aerobic dive duration in marine mammals: from biochemistry to behavior

Marine mammals exhibit multi-level adaptations, from cellular biochemistry to behavior, that maximize aerobic dive duration. A dive response during aerobic dives enables the efficient use of blood and muscle oxygen stores, but it is exercise modulated to maximize the aerobic dive limit at different levels of exertion. Blood volume and concentrations of blood hemoglobin and muscle myoglobin are elevated and serve as a significant oxygen store that increases aerobic dive duration. However, myoglobin is not homogeneously distributed in the locomotory muscles and is highest in areas that produce greater force and consume more oxygen during aerobic swimming. Muscle fibers are primarily fast and slow twitch oxidative with elevated mitochondrial volume densities and enhanced oxidative enzyme activities that are highest in areas that produce more force generation. Most of the muscle mitochondria are interfibriller and homogeneously distributed. This reduces the diffusion distance between mitochondria and helps maintain aerobic metabolism under hypoxic conditions. Mitochondrial volume densities and oxidative enzyme activities are also elevated in certain organs such as liver, kidneys, and stomach. Hepatic and renal function along with digestion and assimilation continue during aerobic dives to maintain physiological homeostasis. Most ATP production comes from aerobic fat metabolism in carnivorous marine mammals. Glucose is derived mostly from gluconeogenesis and is conserved for tissues such as red blood cells and the central nervous system. Marine mammals minimize the energetic cost of swimming and diving through body streamlining, efficient, lift-based propulsive appendages, and cost-efficient modes of locomotion that reduce drag and take advantage of changes in buoyancy with depth. Most dives are within the animal’s aerobic dive limit, which maximizes time underwater and minimizes recovery time at the surface. The result of these adaptations is increased breath-hold duration and enhanced foraging ability that maximizes energy intake and minimizes energy output while making aerobic dives to depth. These adaptations are the long, evolutionary legacy of an aquatic lifestyle that directly affects the fitness of marine mammal species for different diving abilities and environments.     

Diving behaviour of guillemot Uria aalge, puffin Fratercula arctica and razorbill Alca tor da as shown by radio-telemetry

Information on dive and pause times and the numbers of dives in a sequence were obtained for six guillemots and single razorbill and puffin. There were marked differences in diving performance between the species with the order of ranking, in descending order of dive and pause duration, being guillemot, razorbill and puffin. For guillemots, 80% of dives were of 20–119 sec duration and 80% of pauses were 0–59 sec; the maximum dive lasted 202 sec. Puffin dives and pauses were much shorter, with 81% of dives lasting 0–39 sec and 95% of pauses being less than 20 sec, the longest dive was 115 sec. Comparisons of diving sequences made by the same individual indicated some flexibility in all aspects of the sequence but there were broad interspecific differences in the organization of the sequence. The puffin generally made a large number of relatively short dives separated by very short pauses which resulted in a high diving rate (1–5 dives/min) and the bird spending 78% of its time underwater. In contrast, guillemots had much shorter sequences with a few long dives and pauses and lower rates of diving (0–5-0-6 dives/min) and proportions of time underwater (61–65%). Guillemots and puffins may forage at different depths and have different foraging strategies.     

DIVING BEHAVIOR OF THE SAIMAA RINGED SEAL (PHOCA HISPIDA SAIMENSIS NORDQ.)

The activity and diving patterns of four adult Saimaa ringed seals ( Phoca hispida saimensis , a landlocked subspecies living in Lake Saimaa, Finland) were examined during spring, summer, and autumn by the use of VHF-transmitters. Over 17,000 dives were registered. The duration of the dives and diving patterns differed among individuals. The mean duration of dives increased from spring to autumn; e.g. , in one individual the mean dive duration increased from 6 min in June to 10.5 min in October. The haul-out periods of one individual in May to early June made up 46.2% of its total activity budget, but in another individual in July to August the haul-out periods made up only 11% of the budget and the seal was submerged for 80% of the time. Periods of successive long duration dives (>10 min) were observed in three individuals in summer and autumn. The longest dive measured was 23 min. The duration of the periods containing long dives was often over three hours (maximum six hours) and the mean duration of the dives about 15 min. These long duration dives are assumed to be aerobic resting dives. Generally, the dives of the Saimaa ringed seal appear to be of longer duration than previously assumed.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

Diving behaviour, dive cycles and aerobic dive limit in the platypus Ornithorhynchus anatinus

We investigated the diving behaviour, the time allocation of the dive cycle and the behavioural aerobic dive limit (ADL) of platypuses (Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Individual platypuses were equipped with combined data logger-transmitter packages measuring dive depth. Mean dive duration was 31.3 s with 72% of all dives lasting between 18 and 40 s. Mean surface duration was 10.1 s. Mean dive depth was 1.28 m with a maximum of 8.77 m. Platypuses performed up to 1600 dives per foraging trip with a mean of 75 dives per hour. ADL was estimated by consideration of post-dive surface intervals vs. dive durations. Only 15% of all dives were found to exceed the estimated ADL of 40 s, indicating mainly aerobic diving in the species. Foraging platypuses followed a model of optimised recovery time, the optimal breathing theory. Total bottom duration or total foraging duration per day is proposed as a useful indicator of foraging efficiency and hence habitat quality in the species.     

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1+/-0.2 degrees C (n=101 dives in five birds), pectoral muscle: 37.8+/-0.1 degrees C (n=71 dives in three birds) and axillary/brachial veins: 37.9+/-0.1 degrees C (n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=-0.59, P<0.05; range <1 degrees C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6+/-0.7 degrees C (n=157 dives in three birds), 20.2+/-1.2 degrees C (n=69 in three birds) and 35.2+/-0.2 degrees C (n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r=-0.29 to -0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r=-0.49 and -0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 degrees C during rest periods to a grand mean of 15.0+/-0.7 degrees C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r=-0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degrees C more closely than the anterior abdominal temperatures (19-30 degrees C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.     

Sensitivity to hypercapnia and elimination of CO2 following diving in Steller sea lions (Eumetopias jubatus)

The diving ability of marine mammals is a function of how they use and store oxygen and the physiological control of ventilation, which is in turn dependent on the accumulation of CO₂. To assess the influence of CO₂ on physiological control of dive behaviour, we tested how increasing levels of inspired CO₂ (hypercarbia) and decreasing inspired O₂ (hypoxia) affected the diving metabolic rate, submergence times, and dive recovery times (time to replenish O₂ stores and eliminate CO₂) of freely diving Steller sea lions. We also measured changes in breathing frequency of diving and non-diving individuals. Our findings show that hypercarbia increased breathing frequency (as low as 2 % CO₂), but did not affect metabolic rate, or the duration of dives or surface intervals (up to 3 % CO₂). Changes in breathing rates indicated respiratory drive was altered by hypercarbia at rest, but blood CO₂ levels remained below the threshold that would alter normal dive behaviour. It took the sea lions longer to remove accumulated CO₂ than it did for them to replenish their O₂ stores following dives (whether breathing ambient air, hypercarbia, or hypoxia). This difference between O₂ and CO₂ recovery times grew with increasing dive durations, increasing hypercarbia, and was greater for bout dives, suggesting there could be a build-up of CO₂ load with repeated dives. Although we saw no evidence of CO₂ limiting dive behaviour, the longer time required to remove CO₂ may eventually exhibit control over the overall time they can spend in apnoea and overall foraging duration.     

REDUCTIONS IN OXYGEN CONSUMPTION DURING DIVES AND ESTIMATED SUBMERGENCE LIMITATIONS OF STELLER SEA LIONS (EUMETOPIAS JUBATUS)

Accurate estimates of diving metabolic rate are central to assessing the energy needs of marine mammals. To circumvent some of the limitations inherent with conducting energy studies in both the wild and captivity, we measured diving oxygen consumption of two trained Steller sea lions ( Eumetopias jubatus ) in the open ocean. The animals dived to predetermined depths (5–30 m) for controlled periods of time (50–200 s). Rates of oxygen consumption were measured using open-circuit respirometry before and after each dive. Mean resting rates of oxygen consumption prior to the dives were 1.34 (±0.18) and 1.95 (±0.19) liter/min for individual sea lions. Mean rates of oxygen consumption during the dives were 0.71 (±0.24) and 1.10 (±0.39) liter/min, respectively. Overall, rates of oxygen consumption during dives were significantly lower (45% and 41%) than the corresponding rates measured before dives. These results provide the first estimates of diving oxygen consumption rate for Steller sea lions and show that this species can exhibit a marked decrease in oxygen consumption relative to surface rates while submerged. This has important consequences in the evaluation of physiological limitations associated with diving such as dive duration and subsequent interpretations of diving behavior in the wild.