Breathing hypoxic gas affects the physiology as well as the diving behaviour of tufted ducks

We measured the effects of exposure to hypoxia (15% and 11% oxygen) and hypercapnia (up to 4.5% carbon dioxide) on rates of respiratory gas exchange both between and during dives in tufted ducks, Aythya fuligula, to investigate to what extent these may explain changes in diving behaviour. As found in previous studies, the ducks decreased dive duration (t(d)) and increased surface duration when diving from a hypoxic or hypercapnic gas mix. In the hypercapnic conditions, oxygen consumption during the dive cycle was not affected. Oxygen uptake between dives was reduced by only 17% when breathing a hypoxic gas mix of 11% oxygen. However, estimates of the rate of oxygen metabolism during the foraging periods of dives decreased nearly threefold in 11% oxygen. Given that tufted ducks normally dive well within their aerobic dive limits and that they significantly reduced their t(d) during hypoxia, it is not at all clear why they make this physiological adjustment.     

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.     

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.     

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.     

Heart rates and gas exchange in the Amazonian Manatee (Trichechus inunguis) in relation to diving

Unrestrained Amazonian manatees (Trichechus inunguis) maintained a constant heart rate during diving and exhibited a slight tachycardia during breathing. 'Forcing' the manatees to dive caused a marked bradycardia. They exhibited a more pronounced tachycardia during breathing after 'forced' dives and hyperventilated during recovery dives. Manatees are capable of dives exceeding 10 min duration without having to resport to anaerobic metabolism, and even after 10 min dives recover within 3-4 short dives. The ability of manatees to make long dives, in spite of relatively poor O2 stores, is due to their low metabolic rate, while the rapid recovery is aided by their high CO2 stores which minimizes CO2 storage in the body. In manatees the changes in alveolar O2 and CO2 pressure (PAO2 and PACO2) in relation to dive time are slower and more variable than in other marine mammals. The lower rate of change is probably due to the manatees' reduced metabolic rate, while the greater variability is due to their breathing pattern, in which both ventilation and body gas stores influence alveolar gases.     

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.     

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.      

Arterial blood pressure during voluntary diving in the tufted duck,Aythya fuligula

Arterial blood pressure was monitored in voluntarily diving tufted ducks. Mean arterial blood pressure while diving increased during the pre-dive tachycardia, fell to resting levels on submersion, then gradually increased before peaking on surfacing. Estimated total peripheral resistance fell during the pre-dive and post-dive tachycardia, presumably to allow the oxygen stores to be loaded and replenished respectively and/or for carbon dioxide levels to be reduced. Changes in mean arterial blood pressure and total peripheral resistance suggest that peripheral vasoconstriction occurs in some vascular beds during a dive. An increase in arterial blood pressure (and therefore perfusion pressure) may be employed to increase blood flow and oxygen delivery to the active leg muscles.Abbreviations ecg Electrocardiogram, f _H, heart rate - MABP mean arterial blood pressure - P _b blood pressure(s) - TPR total peripheral resistance - V _b cardiac output     

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.     

Aerobic dive limit. What is it and is it always used appropriately?

The original definition of aerobic dive limit (ADL) was the dive duration after which there is an increase in post-dive concentration of lactate in the blood of Weddell seals freely diving in the field. The only other species in which such measurements have been made is the emperor penguin. For all other species, aerobic dive limit has been calculated (cADL) by dividing usable oxygen stores with an estimation of the rate of oxygen consumption during diving. Unfortunately, cADL is often referred to as the aerobic dive limit, implying that it is equivalent to that determined from the measurement of post-dive blood lactate concentration. However, this is not so, as at cADL all of the usable oxygen would have been consumed, whereas Weddell seals and emperor penguins can dive for at least 2-3 times longer than their ADL. Thus, at ADL, there is still some usable oxygen remaining in the stores. It is suggested that to avoid continued confusion between these two terms, the former is called diving lactate threshold (DLT), as it is somewhat analogous to the lactate threshold in exercising terrestrial vertebrates. Possible explanations of how some species routinely dive beyond their cADL are also discussed.     

Role of substance P in hypercapnic excitation of carotid chemoreceptors

Experiments were performed on 17 anesthetized, paralyzed, and artificially ventilated cats to evaluate the importance of substance P-like peptide (SP) on the carotid body responses to CO2. Single or paucifiber carotid chemoreceptor activity was recorded from the peripheral end of the cut carotid sinus nerve. In eight of the cats the influence of SP on hyperoxic hypercapnic responses was studied. While the animals breathed 100% O2, intracarotid infusion of SP (1 microgram.kg-1.min-1, 3 min) increased chemoreceptor activity by +4.8 +/- 0.3 impulses/s. After SP infusion, inhalation of CO2 in O2 caused a rapid increase in activity that reached a peak and then adapted to a lower level, whereas similar levels of CO2 before SP caused only a gradual increase in carotid body discharge rate without any overshoot in response. Furthermore SP significantly increased the magnitude and slope of the CO2 response. In the other nine cats the effect of intracarotid infusion of an SP antagonist, [D-Pro2,D-Trp7,9] SP (10-15 micrograms.kg-1.min-1), on carotid body responses to 1) hyperoxic hypercapnia (7% CO2-93% O2), 2) isocapnic hypoxia (11% O2-89% N2), and 3) hypoxic hypercapnia (11% O2-7% CO2-82% N2) was examined. SP antagonist had no effect on carotid body response to hyperoxic hypercapnia but significantly attenuated the chemoreceptor excitation caused by isocapnic hypoxia and hypoxic hypercapnia. These results suggest that 1) SP may play an important role in carotid body responses to hypoxia but not to CO2, and 2) the mechanisms of stimulation of the carotid body by hypercapnia and by hypoxia differ.     

Estimating the rate of oxygen consumption during submersion from the heart rate of diving animals

How animals manage their oxygen stores during diving and other breath-hold activities has been a topic of debate among physiologists for decades. Specifically, while the behavior of free-ranging diving animals suggests that metabolism during submersion must be primarily aerobic in nature, no studies have been able to determine their rate of oxygen consumption during submersion (Vo(2)d) and hence prove that this is the case. In the present study, we combine two previously used techniques and develop a new model to estimate Vo(2)d accurately and plausibly in a free-ranging animal and apply it to data for macaroni penguins (Eudyptes chrysolophus) as an example. For macaroni penguins at least, Vo(2)d can be predicted by measuring heart rate during the dive cycle and the subsequent surface interval duration. Including maximum depth of the dive improves the accuracy of these predictions. This suggests that energetically demanding locomotion events within the dive combine with the differing buoyancy and locomotion costs associated with traveling to depth to influence its cost in terms of oxygen use. This will in turn effect the duration of the dive and the duration of the subsequent recovery period. In the present study, Vo(2)d ranged from 4 to 28 ml.min(-1).kg(-1), indicating that, at least as far as aerobic metabolism was concerned, macaroni penguins were often hypometabolic, with rates of oxygen consumption usually below that for this species resting in water (25.6 ml.min(-1).kg(-1)) and occasionally lower than that while resting in air (10.3 ml.min(-1).kg(-1)).     

Pattern of the ventilatory response to transient hypoxia in man: differences from transient hypercapnic test.

The pattern of change in ventilatory variables after inhalation of pure N2 for two breaths was studied in normal children and adults. In six subjects the trends of change were compared to the ventilatory response to transient hypercapnia. We observed differences in the patterns of increasing ventilation with an initial abrupt increase of tidal volume for transient hypoxia and a progressive change for hypercapnia. In both cases respiratory frequency was progressively but unsystematically enhanced. A highly significant positive correlation was demonstrated between individual sensitivities to CO2 and O2, with a greater response to hypercapnia (5.6 time) than to hypoxia. Finally, a very short-latency decrease in expiratory duration occurred in the first breath after inhalation of hypercapnic mixture, supporting the recent data of Cunningham et al. (1977).     

Physiological and behavioral determinants of the aerobic dive limit in Weddell seal (Leptonychotes weddellii) Pups

The aerobic dive limit, as defined by an increase in plasma lactate levels following dives, has to date only been determined in adult and juvenile Weddell seals (Leptonychotes weddellii). However, theoretical aerobic dive limits based on calculated total body oxygen stores, estimated metabolic rates, and dive duration frequencies have been published for several species. Using data collected over the past 3 years in McMurdo Sound. Antarctica, the aerobic dive limit of Weddell seal pups was determined by both the physiological and modeling methods. Time-depth diving recorders deployed on 36 pups between 2 and 14 weeks of age allowed the aerobic dive limit to be predicted from duration-frequency histograms. The aerobic dive limit was also calculated from estimates of total body oxygen stores and predicted diving metabolic rates. Finally, these two estimates were compared with aerobic dive limits determined from post-dive lactate levels in three pups between 5 and 7 weeks old. The aerobic dive limits of pups increased with age, but pup aerobic dive limits were still significantly shorter than those of yearlings and adults. In addition, the aerobic dive limits determined by the three methods were not equivalent for pups, yearlings, or adults, and indicate that care should be taken when modeling methods are used to estimate the aerobic dive limit in other species. Changes in hematocrit, plasma glucose, and plasma lactate levels during and between rest, diving, and recovery in pups were compared to known values for juveniles and adults. Plasma metabolite levels were more highly regulated in older pups, and together with the increasing aerobic dive limit, suggest that Weddell seal pups are not refined divers until after they are weaned, and that their diving ability continues to develop over several years.     

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.     

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.     

Investigating Annual Diving Behaviour by Hooded Seals (Cystophora cristata) within the Northwest Atlantic Ocean

With the exception of relatively brief periods when they reproduce and moult, hooded seals, Cystophora cristata, spend most of the year in the open ocean where they undergo feeding migrations to either recover or prepare for the next fasting period. Valuable insights into habitat use and diving behaviour during these periods have been obtained by attaching Satellite Relay Data Loggers (SRDLs) to 51 Northwest (NW) Atlantic hooded seals (33 females and 18 males) during ice-bound fasting periods (2004−2008). Using General Additive Models (GAMs) we describe habitat use in terms of First Passage Time (FPT) and analyse how bathymetry, seasonality and FPT influence the hooded seals’ diving behaviour described by maximum dive depth, dive duration and surface duration. Adult NW Atlantic hooded seals exhibit a change in diving activity in areas where they spend >20 h by increasing maximum dive depth, dive duration and surface duration, indicating a restricted search behaviour. We found that male and female hooded seals are spatially segregated and that diving behaviour varies between sexes in relation to habitat properties and seasonality. Migration periods are described by increased dive duration for both sexes with a peak in May, October and January. Males demonstrated an increase in dive depth and dive duration towards May (post-breeding/pre-moult) and August–October (post-moult/pre-breeding) but did not show any pronounced increase in surface duration. Females dived deepest and had the highest surface duration between December and January (post-moult/pre-breeding). Our results suggest that the smaller females may have a greater need to recover from dives than that of the larger males. Horizontal segregation could have evolved as a result of a resource partitioning strategy to avoid sexual competition or that the energy requirements of males and females are different due to different energy expenditure during fasting periods.     

A theoretical analysis of diving performance in the Weddell seal (Leptonychotes weddelli)

Marine mammals are constrained in their foraging behaviour because, as obligate air breathers, they must undertake regular trips to the water surface to satisfy the need for respiratory gas exchange. Maximum underwater endurance time is determined by O2 supply and demand, but this does not necessarily imply that O2 is the main factor regulating individual dive and surface times. This study presents a theoretical analysis of diving performance that emphasizes a key role for CO2 in the proximate control of diving behaviour. Computer simulations, based on a mathematical model of the mammalian cardiorespiratory control system, are used to investigate the influence of swimming to depth and other energetic stresses (feeding, thermogenesis, sleep) on predicted diving behaviour in an average adult Weddell seal. The plausibility of the proposed model is supported by the study, which replicated published observations of natural diving behaviour in this species. It is suggested that diving behaviour is tuned to oscillations in respiratory drive and that behavioural and physiological factors can alter the dynamic characteristics of the system to achieve a highly adaptable reciprocal interaction that blurs the boundary between physiology and behaviour.     

Foraging by deep-diving birds is not constrained by an aerobic diving limit: a model of avian depth-dependent diving metabolic rate

The theoretical aerobic diving limit (tADL) specifies the duration of a dive after which oxygen reserves available for diving are depleted. The tADL has been calculated by dividing the available oxygen stores by the diving metabolic rate (DMR). Contrary to diving mammals, most diving birds examined to date exceed the tADL by a large margin. This discrepancy between observation and theory has engendered two alternative explanations suggesting that dive duration is extended either anaerobically or by depressing aerobic metabolism. Current formulations of tADL uncritically assume that DMR is independent of depth. However, diving birds differ from other vertebrate divers by having a larger respiratory system volume and by retaining air in their plumage while diving, thereby elevating buoyancy. Because air compresses with depth, diving power requirement decreases with depth. Following this principle, we modeled DMR to depth for Adelie and little penguins and reformulated the tADL accordingly. The model's results suggest that < approximately 5% of natural dives by Adelie penguins exceed the reformulated tADL(d), or maximal aerobic depth, and none in the more buoyant little penguin. These data suggest that, for both small and large species, deep diving birds rarely if ever exceed tADL(d).     

High diving metabolism results in a short aerobic dive limit for Steller sea lions (Eumetopias jubatus)

The diving capacity of marine mammals is typically defined by the aerobic dive limit (ADL) which, in lieu of direct measurements, can be calculated (cADL) from total body oxygen stores (TBO) and diving metabolic rate (DMR). To estimate cADL, we measured blood oxygen stores, and combined this with diving oxygen consumption rates (VO₂) recorded from 4 trained Steller sea lions diving in the open ocean to depths of 10 or 40 m. We also examined the effect of diving exercise on O₂ stores by comparing blood O₂ stores of our diving animals to non-diving individuals at an aquarium. Mass-specific blood volume of the non-diving individuals was higher in the winter than in summer, but there was no overall difference in blood O₂ stores between the diving and non-diving groups. Estimated TBO (35.9 ml O₂ kg^?1) was slightly lower than previously reported for Steller sea lions and other Otariids. Calculated ADL was 3.0 min (based on an average DMR of 2.24 L O₂ min^?1) and was significantly shorter than the average 4.4 min dives our study animals performed when making single long dives—but was similar to the times recorded during diving bouts (a series of 4 dives followed by a recovery period on the surface), as well as the dive times of wild animals. Our study is the first to estimate cADL based on direct measures of VO₂ and blood oxygen stores for an Otariid and indicates they have a much shorter ADL than previously thought.