Search

Comparative physiology has developed a rich understanding of the physiological adaptations of organisms, from microbes to megafauna. Despite extreme differences in size and a diversity of habitats, general patterns are observed in their physiological adaptations. Yet, many organisms deviate from the general patterns, providing an opportunity to understand the importance of ecology in determining the evolution of unusual adaptations. Aquatic air-breathing vertebrates provide unique study systems in which the interplay between ecology, physiology and behavior is most evident. They must perform breath-hold dives to obtain food underwater, which imposes a physiological constraint on their foraging time as they must resurface to breathe. This separation of two critical resources has led researchers to investigate these organisms' physiological adaptations and trade-offs. Addressing such questions on large marine animals is best done in the field, given the difficulty of replicating the environment of these animals in the lab. This Review examines the long history of research on diving physiology and behavior. We show how innovative technology and the careful selection of research animals have provided a holistic understanding of diving mammals' physiology, behavior and ecology. We explore the role of the aerobic diving limit, body size, oxygen stores, prey distribution and metabolism. We then identify gaps in our knowledge and suggest areas for future research, pointing out how this research will help conserve these unique animals., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Survival of air-breathing, diving ectotherms is dependent on their capacity to optimise the time available for obligate underwater activities, such as predator avoidance. Submergence times are thermally sensitive, with dive durations significantly reduced by increases in water temperature, deeming these animals particularly vulnerable to the effects of climate change. The physiological mechanisms underlying this compromised performance are unclear but are hypothesised to be linked to increased oxygen demand and a reduced capacity for metabolic depression at elevated temperatures. Here, we investigated how water temperature (both acute and chronic exposures) affected the physiology of juvenile estuarine crocodiles (Crocodylus porosus) performing predator avoidance dives (i.e. fright-dives). Diving oxygen consumption, 'fright' bradycardia, haematocrit and haemoglobin (indicators of blood oxygen carrying capacity) were assessed at two test temperatures, reflective of different climate change scenarios (i.e. current summer water temperatures, 28°C, and 'high' climate warming, 34°C). Diving oxygen consumption rate increased threefold between 28 and 34°C (Q10=7.4). The capacity to depress oxygen demand was reduced at elevated temperatures, with animals lowering oxygen demand from surface levels by 52.9±27.8% and 27.8±16.5% (means±s.e.m.) at 28°C and 34°C, respectively. Resting and post-fright-dive haematocrit and haemoglobin were thermally insensitive. Together these findings suggest decrements in fright-dive performance at elevated temperatures stem from increased oxygen demand coupled with a reduced capacity for metabolic depression. [ABSTRACT FROM AUTHOR]

Among pinnipeds, southern elephant seals (SESs, Mirounga leonina) are extreme divers that dive deeply and continuously along foraging trips to restore their body stores after fasting on land during breeding or moulting. Their replenishment of body stores influences their energy expenditure during dives and their oxygen (O2) reserves (via muscular mass), yet how they manage their O2 stores during their dives is not fully understood. In this study, 63 female SESs from Kerguelen Island were equipped with accelerometers and time-depth recorders to investigate changes in diving parameters through their foraging trips. Two categories of dive behaviour were identified and related to the body size of individuals, with smaller SESs performing shallower and shorter dives requiring greater mean stroke amplitude compared with larger individuals. In relation to body size, the larger seals had lower estimated oxygen consumption levels for a given buoyancy (i.e. body density) compared with smaller individuals. However, both groups were estimated to have the same oxygen consumption of 0.079±0.001 ml O2 stroke-1 kg-1 for a given dive duration and at neutral buoyancy when the cost of transport was minimal. Based on these relationships, we built two models that estimate changes in oxygen consumption according to dive duration and body density. The study highlights that replenishing body stores improves SES foraging efficiency, as indicated by increased time spent at the bottom of the ocean. Thus, prey-capture attempts increase as SES buoyancy approaches the neutral buoyancy point., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Our understanding of how air-breathing marine predators cope with environmental variability is limited by our inadequate knowledge of their ecological and physiological parameters. Because of their wide distribution along both coasts of the sub-continent, South American sea lions (Otaria byronia) provide a valuable opportunity to study the behavioral and physiological plasticity of a marine predator in different environments. We measured the oxygen stores and diving behavior of South American sea lions throughout most of its range, allowing us to demonstrate that diving ability and behavior vary across its range. We found no significant differences in mass-specific blood volumes of sea lions among field sites and a negative relationship between mass-specific oxygen storage and size, which suggests that exposure to different habitats and geographical locations better explains oxygen storage capacities and diving capability in South American sea lions than body size alone. The largest animals in our study (individuals from Uruguay) were the shallowest and shortest duration divers, and had the lowest mass-specific total body oxygen stores, while the deepest and longest duration divers (individuals from southern Chile) had significantly larger mass-specific oxygen stores, despite being much smaller animals. Our study suggests that the physiology of air-breathing diving predators is not fixed, but that it can be adjusted, to a certain extent, depending on the ecological setting and or habitat. These adjustments can be thought of as a 'training effect': as the animal continues to push its physiological capacity through greater hypoxic exposure, its breath-holding capacity increases. [ABSTRACT FROM AUTHOR]

Physiological constraints dictate animals' ability to exploit habitats. For marine mammals, it is important to quantify physiological limits that influence diving and their ability to alter foraging behaviors. We characterized age-specific dive limits of walruses by measuring anaerobic (acid-buffering capacity) and aerobic (myoglobin content) capacities of the muscles that power hind (longissimus dorsi) and fore (supraspinatus) flipper propulsion. Mean buffering capacities were similar acrossmuscles and age classes (a fetus, five neonatal calves, a 3 month old and 20 adults), ranging from 41.31 to 54.14 slykes and 42.00 to 46.93 slykes in the longissimus and supraspinatus, respectively. Mean myoglobin in the fetus and neonatal calves fell within a narrow range (longissimus: 0.92-1.68 g 100 g-1 wet muscle mass; supraspinatus: 0.88-1.64 g 100 g-1 wet muscle mass). By 3 monthspost-partum,myoglobin in the longissimus increased by79%, but levels in the supraspinatus remained unaltered. From 3 months post-partum to adulthood, myoglobin increased by an additional 26% in the longissimus and increased by 126% in the supraspinatus; myoglobin remained greater in the longissimus compared with the supraspinatus. Walruses are unique among marine mammals because theyare born withamaturemuscleacid-buffering capacityand attainmaturemyoglobin content early in life.Despite rapid physiological development, small body size limits the diving capacity of immature walruses and extreme sexual dimorphism reduces the diving capacity of adult females compared with adult males. Thus, free-ranging immature walruses likely exhibit the shortest foraging dives while adult males are capable of the longest foraging dives. [ABSTRACT FROM AUTHOR]

Myoglobin (Mb) isanoxygenbinding protein found in vertebrate skeletal muscle, where it facilitates intracellular transport and storage of oxygen. This protein has evolved to suit unique physiological needs in the muscle of diving vertebrates that express Mb at much greater concentrations than their terrestrial counterparts. In this study, we characterized Mb oxygen affinity (P50) from 25 species of aquatic and terrestrial birds and mammals. Among diving species, we tested for correlations between Mb P50and routine dive duration. Across all species examined, Mb P50ranged from 2.40 to 4.85 mmHg. Themean P50ofMbfromterrestrial ungulateswas 3.72±0.15 mmHg(range 3.70- 3.74 mmHg). The P50of cetaceans was similar to terrestrial ungulates ranging from 3.54 to 3.82 mmHg, with the exception of the melonheaded whale, which had a significantly higher P50of 4.85 mmHg. Among pinnipeds, the P50ranged from3.23 to 3.81 mmHgand showed a trend for higher oxygen affinity in species with longer dive durations. Among diving birds, the P50ranged from 2.40 to 3.36 mmHg and also showed a trend of higher affinities in species with longer dive durations. In pinnipeds and birds, lowMb P50was associated with species whose muscles are metabolically active under hypoxic conditions associated with aerobic dives. Given the broad range of potential globin oxygen affinities, Mb P50from diverse vertebrate species appears constrained within a relatively narrow range. High Mb oxygen affinity within this rangemay be adaptive for some vertebrates thatmake prolonged dives. [ABSTRACT FROM AUTHOR]

Most marine mammals are hypothesized to routinely dive within their aerobic dive limit (ADL). Mammals that regularly perform deep, long-duration dives have locomotor muscles with elevated myoglobin concentrations that are composed of predominantly large, slow-twitch (Type I) fibers with low mitochondrial volume densities (Vmt). These features contribute to extending ADL by increasing oxygen stores and decreasing metabolic rate. Recent tagging studies, however, have challenged the view that two groups of extreme deep-diving cetaceans dive within their ADLs. Beaked whales (including Ziphius cavirostris and Mesoplodon densirostris) routinely perform the deepest and longest average dives of any air-breathing vertebrate, and short-finned pilot whales (Globicephala macrorhynchus) perform high-speed sprints at depth. We investigated the locomotor muscle morphology and estimated total body oxygen stores of several species within these two groups of cetaceans to determine whether they (1) shared muscle design features with other deep divers and (2) performed dives within their calculated ADLs. Muscle of both cetaceans displayed high myoglobin concentrations and large fibers, as predicted, but novel fiber profiles for diving mammals. Beaked whales possessed a sprinter's fiber-type profile, composed of ~80% fast-twitch (Type II) fibers with low VmX. Approximately one-third of the muscle fibers of short-finned pilot whales were slow-twitch, oxidative, glycolytic fibers, a rare fiber type for any mammal. The muscle morphology of beaked whales likely decreases the energetic cost of diving, while that of short-finned pilot whales supports high activity events. Calculated ADLs indicate that, at low metabolic rates, both beaked and short-finned pilot whales carry sufficient onboard oxygen to aerobically support their dives. [ABSTRACT FROM AUTHOR]

Since the introduction of the aerobic dive limit (ADL) 30 years ago, the concept that most dives of marine mammals and sea birds are aerobic in nature has dominated the interpretation of their diving behavior and foraging ecology. Although there have been many measurements of body oxygen stores, there have been few investigations of the actual depletion of those stores during dives. Yet, it is the pattern, rate and magnitude of depletion of O2 stores that underlie the ADL. Therefore, in order to assess strategies of O2 store management, we review (a) the magnitude of 02 stores, (b) past studies of O2 store depletion and (C) our recent investigations of O2 store utilization during sleep apnea and dives of elephant seals (Mirounga angustirostris) and during dives of emperor penguins (Aptenodytes forsteri). We conclude with the implications of these findings for (a) the physiological responses underlying O2 store utilization, (b) the physiological basis of the ADL and (c) the value of extreme hypoxemic tolerance and the significance of the avoidance of re-perfusion injury in these animals. [ABSTRACT FROM AUTHOR]

Emperor penguins (Aptenodytes forsteri), both at sea and at an experimental dive hole, often have minimal surface periods even after performance of dives far beyond their measured 5.6 min aerobic dive limit (ADL: dive duration associated with the onset of post-dive blood lactate accumulation). Accelerometer-based data loggers were attached to emperor penguins diving in these two different situations to further evaluate the capacity of these birds to perform such dives without any apparent prolonged recovery periods. Minimum surface intervals for dives as long as 10 min were less than 1 min at both sites. Stroke rates for dives at sea were significantly greater than those for dives at the isolated dive hole. Calculated diving air volumes at sea were variable, increased with maximum depth of dive to a depth of 250 m, and decreased for deeper dives. It is hypothesized that lower air volumes for the deepest dives are the result of exhalation of air underwater. Mean maximal air volumes for deep dives at sea were approximately 83% greater than those during shallow (<50 m) dives. We conclude that (a) dives beyond the 5.6 min ADL do not always require prolongation of surface intervals in emperor penguins, (b) stroke rate at sea is greater than at the isolated dive hole and, therefore, a reduction in muscle stroke rate does not extend the duration of aerobic metabolism during dives at sea, and (c) a larger diving air volume facilitates performance of deep dives by increasing the total body O2 store to 68 ml O2 kg-1. Although increased O2 storage and cardiovascular adjustments presumably optimize aerobic metabolism during dives, enhanced anaerobic capacity and hypoxemic tolerance are also essential for longer dives. This was exemplified by a 27.6 min dive, after which the bird required 6 min before it stood up from a prone position, another 20 min before it began to walk, and 8.4 h before it dived again. [ABSTRACT FROM AUTHOR]

The physiological basis of the aerobic dive limit (ADL), the dive duration associated with the onset of post-dive blood lactate elevation, is hypothesized to be depletion of the muscle oxygen (O2) store. A duel wavelength near-infrared spectrophotometer was developed and used to measure myoglobin (Mb) O2 saturation levels in the locomotory muscle during dives of emperor penguins (Aptenodytes forsteri). Two distinct patterns of muscle O2 depletion were observed. Type A dives had a monotonic decline, and, in dives near the ADL, the muscle O2 store was almost completely depleted. This pattern of Mb desaturation was consistent with lack of muscle blood flow and supports the hypothesis that the onset of post-dive blood lactate accumulation is secondary to muscle O2 depletion during dives. The mean type A Mb desaturation rate allowed for calculation of a mean muscle O2 consumption of 12.4 ml O2kg-1 muscle min-1, based on a Mb concentration of 6.4g 100g-1 muscle. Type B desaturation patterns demonstrated a more gradual decline, often reaching a mid-dive plateau in Mb desaturation. This mid-dive plateau suggests maintenance of some muscle perfusion during these dives. At the end of type B dives, Mb desaturation rate increased and, in dives beyond the ADL, Mb saturation often reached near 0%. Thus, although different physiological strategies may be used during emperor penguin diving, both Mb desaturation patterns support the hypothesis that the onset of post-dive lactate accumulation is secondary to muscle O2 store depletion [ABSTRACT FROM AUTHOR]

Marine turtles spend more than 90% of their life underwater and have been termed surfacers as opposed to divers. Nonetheless turtles have been reported occasionally to float motionless at the surface but the reasons for this behaviour are not clear. We investigated the location, timing and duration of extended surface times (ESTs) in 10 free-ranging loggerhead turtles (Caretta caretta) and the possible relationship to water temperature and diving activity recorded via satellite relay data loggers for 101-450 days. For one turtle that dived only in offshore areas, ESTs contributed 12% of the time whereas for the other turtles ESTs contributed 0.4-1.8% of the time. ESTs lasted on average 90 mm but were mostly infrequent and irregular, excluding the involvement of a fundamental regulatory function. However, 82% of the ESTs occurred during daylight, mostly around noon, suggesting a dependence on solar radiation. For three turtles, there was an appreciable (7°C to 10.5°C) temperature decrease with depth for dives during periods when ESTs occurred frequently, suggesting a re-warming function of EST to compensate for decreased body temperatures, possibly to enhance digestive efficiency. A positive correlation between body mass and EST duration supported this explanation. By contrast, night-active turtles that exceeded their calculated aerobic dive limits in 7.6-16% of the dives engaged in nocturnal ESTs, probably for lactate clearance. This is the first evidence that loggerhead turtles may refrain from diving for at least two reasons, either to absorb solar radiation or to recover from anaerobic activity. [ABSTRACT FROM AUTHOR]

The emperor penguin (Aptenodytes forsteri) thrives in the Antarctic underwater environment, diving to depths greater than 500m and for durations longer than 23 mm. To examine mechanisms underlying the exceptional diving ability of this species and further describe blood oxygen (O2) transport and depletion while diving, we characterized the O2-hemoglobin (Hb) dissociation curve of the emperor penguin in whole blood. This allowed us to (1) investigate the biochemical adaptation of Hb in this species, and (2) address blood O2 depletion during diving, by applying the dissociation curve to previously collected partial pressure of O2 (PO2) profiles to estimate in vivo Hb saturation (SO2) changes during dives. This investigation revealed enhanced Hb-O2 affinity (P50=28 mmHg, pH7.5) in the emperor penguin, similar to high-altitude birds and other penguin species. This allows for increased O2 at low blood PO2 levels during diving and more complete depletion of the respiratory O2 store. SO2 profiles during diving demonstrated that arterial 8O2 levels are maintained near 100% throughout much of the dive, not decreasing significantly until the final ascent phase. End-of-dive venous SO2 values were widely distributed and optimization of the venous blood O2 store resulted from arterialization and near complete depletion of venous blood O2 during longer dives. The estimated contribution of the blood O2 store to diving metabolic rate was low and highly variable. This pattern is due, in part, to the influx of O2 from the lungs into the blood during diving, and variable rates of tissue O2 uptake. [ABSTRACT FROM AUTHOR]

In order to further define O2 store utilization during dives and understand the physiological basis of the aerobic dive limit (ADL, dive duration associated with the onset of post-dive blood lactate accumulation), emperor penguins (Aptenodytes forsteri) were equipped with either a blood partial pressure of oxygen (PO2) recorder or a blood sampler while they were diving at an Isolated dive hole in the sea ice of McMurdo Sound, Antarctica. Arterial PO2 profiles (57 dives) revealed that (a) pre-dive PO2 was greater than that at rest, (b) PO2 transiently increased during descent and (c) post-dive PO2 reached that at rest in 1.92±1.89 mm (N=53). Venous PO2 profiles (130 dives) revealed that (a) pre-dive venous PO2 was greater than that at rest prior to 61% of dives, (b) in 90% of dives venous PO2 transiently increased with a mean maximum PO2 of 53±18mmHg and a mean increase in PO2 of 11±12mmHg, (c) in 78% of dives, this peak venous PO2 occurred within the first 3mm, and (d) post-dive venous PO2 reached that at rest within 2.23±2.64mm (N=84). Arterial and venous PO2 values in blood samples collected 1-3mm into dives were greater than or near to the respective values at rest. Blood lactate concentration was less than 2 mmoll-1 as far as 10.5 mm into dives, well beyond the known ADL of 5.6mm. Mean arterial and venous of samples collected at 20-37m depth were 2.5 times those at the surface, both being 2.1 ±0.7 atmospheres absolute (ATA;N=3 each), and were not significantly different. These findings are consistent with the maintenance of gas exchange during dives (elevated arterial and venous PO2 and PN2 during dives), muscle ischemia during dives (elevated venous PO2, lack of lactate washout into blood during dives), and arterio-venous shunting of blood both during the surface period (venous PO2 greater than that at rest) and during dives (arterialized venous PO2 values during descent, equivalent arterial and venous PN2 values during dives). These three physiological processes contribute to the transfer of the large respiratory O2 store to the blood during the dive, isolation of muscle metabolism from the circulation during the dive, a decreased rate of blood O2 depletion during dives, and optimized loading of O2 stores both before and after dives. The lack of blood O2 depletion and blood lactate elevation during dives beyond the AOL suggests that active locomotory muscle is the site of tissue lactate accumulation that results in post-dive blood lactate elevation in dives beyond the AOL. [ABSTRACT FROM AUTHOR]

Blood gas analyses from emperor penguins (Aptenodytes forsteri) at rest, and intravascular PO2 profiles from free- diving birds were obtained in order to examine hypoxemic tolerance and utilization of the blood O2 store during dives. Analysis of blood samples from penguins at rest revealed arterial PO2s and O2 contents of 68±7 mmHg (1 mmHg= 133.3 Pa) and 22.5±1.3 ml O2 dl-1 (N=3) and venous values of 41±10 mmHg and 17.4±2.9 ml O2 dl-1 (N=9). Corresponding arterial and venous Hb saturations for a hemoglobin (Hb) concentration of 18 g dl-1 were >91% and 70%, respectively. Analysis of PO2 profiles obtained from birds equipped with intravascular PO2 electrodes and backpack recorders during dives revealed that (1) the decline of the final blood PO2 of a dive in relation to dive duration was variable, (2) final venous PO2 values spanned a 40-mmHg range at the previously measured aerobic dive limit (ADL; dive duration associated with onset of post-dive blood lactate accumulation), (3) final arterial, venous and previously measured air sac PO2 values were indistinguishable in longer dives, and (4) final venous PO2 values of longer dives were as low as 1-6 mmHg during dives. Although blood O2 is not depleted at the ADL, nearly complete depletion of the blood O2 store occurs in longer dives. This extreme hypoxemic tolerance, which would be catastrophic in many birds and mammals, necessitates biochemical and molecular adaptations, including a shift in the O2-Hb dissociation curve of the emperor penguin in comparison to those of most birds. A relatively higher-affinity Hb is consistent with blood PO2 values and O2 contents of penguins at rest. [ABSTRACT FROM AUTHOR]

A mathematical model was used to explore if elevated levels of N2, and risk of decompression sickness (DCS), could limit dive performance (duration and depth) in king penguins (Aptenodytes patagonicus). The model allowed prediction of blood and tissue (central circulation, muscle, brain and fat) N2 tensions (PN2) based on different cardiac outputs and blood flow distributions. Estimated mixed venous PN2 agreed with values observed during forced dives in a compression chamber used to validate the assumptions of the model. During bouts of foraging dives, estimated mixed venous and tissue PN2 increased as the bout progressed. Estimated mean maximum mixed venous PN2 upon return to the surface after a dive was 4.56±0.18 atmospheres absolute (ATA; range: 4.37–4.78 ATA). This is equivalent to N2 levels causing a 50% DCS incidence in terrestrial animals of similar mass. Bout termination events were not associated with extreme mixed venous N2 levels. Fat PN2 was positively correlated with bout duration and the highest estimated fat PN2 occurred at the end of a dive bout. The model suggested that short and shallow dives occurring between dive bouts help to reduce supersaturation and thereby DCS risk. Furthermore, adipose tissue could also help reduce DCS risk during the first few dives in a bout by functioning as a sink to buffer extreme levels of N2. [ABSTRACT FROM AUTHOR]

A defining physiological capability for air-breathing marine vertebrates is the amount of oxygen that can be stored in tissues and made available during dives. To evaluate the influence of oxygen storage capacity on aerobic diving capacity, we examined the extent to which blood and muscle oxygen stores varied as a function of age, body size and sex in the sexually dimorphic California sea lion, Zalophus californianus. We measured total body oxygen stores, including hematocrit, hemoglobin, MCHC, plasma volume, blood volume and muscle myoglobin in pups through adults of both sexes. Blood and muscle oxygen storage capacity was not fully developed by the end of the dependency period, with blood stores not fully developed until animals were larger juveniles (70 kg; 1.5-2.5 years) and muscle stores not until animals were sub-adult size (125 kg; 4-6 years). Differences in aerobic diving capacity among size classes were reflective of these major milestones in the development of oxygen stores. Male sea lions had greater absolute blood volume than females and reflected the larger mass of males, which became apparent when animals were large juveniles. Adult female sea lions had greater muscle myoglobin concentrations compared to males, resulting in greater mass-specific muscle and total oxygen stores. Delayed development of oxygen stores is consistent with the shallow epi-mesopelagic foraging behavior in this species. We hypothesize that the greater mass-specific oxygen stores of female sea lions compared to males is related to differences in foraging behavior between the sexes. [ABSTRACT FROM AUTHOR]

Sound-and-orientation recording tags (DTAGs) were used to study 10 beaked whales of two poorly known species, Ziphius cavirostris (Zc) and Mesoplodon densirostris (Md). Acoustic behaviour in the deep foraging dives performed by both species (Zc: 28 dives by seven individuals; Md: 16 dives by three individuals) shows that they hunt by echolocation in deep water between 222 and 1885 m, attempting to capture about 30 prey/dive. This food source is so deep that the average foraging dives were deeper (Zc: 1070 m; Md: 835 m) and longer (Zc: 58 mm; Md: 47 mm) than reported for any other air-breathing species. A series of shallower dives, containing no indications of foraging, followed most deep foraging dives. The average interval between deep foraging dives was 63 mm for Zc and 92 mm for Md. This long an interval may be required for beaked whales to recover from an oxygen debt accrued in the deep foraging dives, which last about twice the estimated aerobic dive limit. Recent reports of gas emboli in beaked whales stranded during naval sonar exercises have led to the hypothesis that their deep-diving may make them especially vulnerable to decompression. Using current models of breath-hold diving, we infer that their natural diving behaviour is inconsistent with known problems of acute nitrogen supersaturation and embolism. If the assumptions of these models are correct for beaked whales, then possible decompression problems are more likely to result from an abnormal behavioural response to sonar. [ABSTRACT FROM AUTHOR]

The main objective of this study was to determine heart rate (fH) and the energetic costs of specific behaviours of king penguins while ashore and while foraging at sea during their breeding period. In particular, an estimate was made of the energetic cost of diving in order to determine the proportion of dives that may exceed the calculated aerobic dive limit (cADL; estimated usable O2 stores/estimated rate of oxygen consumption during diving). An implanted data logger enabled fH and diving behaviour to be monitored from 10 free-ranging king penguins during their breeding period. Using previously determined calibration equations, it was possible to estimate rate of oxygen consumption (&Vdot;O2) when the birds were ashore and during various phases of their foraging trips. Diving behaviour showed a clear diurnal pattern, with a mixture of deep (>40 m), long (>3 mm) and shallow (<40 m), short (<3 mm) dives from dawn to dusk and shallow, short dives at night. Heart rate during dive bouts and dive cycles (dive + post-dive interval) was 42% greater than that when the birds were ashore. During diving, fH was similar to the ‘ashore’ value (87±4 beats min-1), but it did decline to 76% of the value recorded from king penguins resting in water. During the first hour after a diving bout, fH was significantly higher than the average value during diving (101±4 beats min-1) and for the remainder of the dive bout. Rates of oxygen consumption estimated from these (and other) values of fit indicate that when at sea, metabolic rate (MR) was 83% greater than that when the birds were ashore [3.15 W kg-1 (-0.71, +0.93), where the values in parentheses are the computed standard errors of the estimate], while during diving bouts and dive cycles, It was 73% greater than the ‘ashore’ value. Although estimated MR during the total period between dive bouts was not significantly different from that during dive bouts [5.44 W kg-1 (-0.30, +0.32)], MR during the first hour following a dive bout was 52% greater than that during a diving bout. It is suggested that this large increase following diving (foraging) activity is, at least in part, the result of rewarming the body, which occurs at the end of a diving bout. From the measured behaviour and estimated values of &Vdot;O2, it was evident that approximately 35% of the dives were in excess of the cADL. Even If &Vdot;O2 during diving was assumed to be the same as when the birds were resting on water, approximately 20% of dives would exceed the cADL. As &Vdot;O2 during diving is, in fact, that estimated for a complete dive cycle, it is quite feasible that &Vdot;O2 during diving itself is less than that measured for birds resting in water, it is suggested that the regional hypothermia that has been recorded in this species during diving bouts may be at least a contributing factor to such hypometabolism. [ABSTRACT FROM AUTHOR]

The energetic cost of diving in marine mammals is a difficult value to derive given the problems of assessing metabolic rate for an animal at sea. Nevertheless, it is fundamental to our understanding of the foraging strategies of air-breathers exploiting underwater food sources. We measured the metabolic rates of eight captive grey seals, voluntarily diving in a quasi-natural setting. Oxygen consumption during post-dive surface periods was measured using open-flow respirometry, and dive behaviour of the seals was recorded using time depth recorders (TDRs). Mean diving metabolic rate (DMR) for both adults and juveniles was 1.7 times the predicted standard metabolic rate of terrestrial animals of equal size. For all animals, DMR was lower than the rate of metabolism measured whilst they were resting at the water's surface. On a dive-by-dive basis, DMR decreased with dive duration but increased with mean swim speed. Regressing the maximum 5% of DMRs against dive duration resulted in a significant negative relationship that was not significantly different from the relationship between the calculated maximum rate of aerobic metabolism and dive duration, suggesting that these seals were diving within, and up to, their aerobic limits. We developed a model that allows the prediction of DMR from information on dive behaviour of the type routinely collected in telemetry studies of wild seals. The model accurately predicts DMR using behavioural data from periods of diving with known metabolism data. This model can be used to predict the at-sea metabolic rate of wild grey seals, an important input into ecosystem models. [ABSTRACT FROM AUTHOR]

Diving ectothermic vertebrates are an important component of many aquatic ecosystems, but the threat of climate warming is particularly salient to this group. Dive durations typically decrease as water temperatures rise; yet, we lack an understanding of whether this trend is apparent in all diving ectotherms and how this group will fare under climate warming. We compiled data from 27 studies on 20 ectothermic vertebrate species to quantify the effect of temperature on dive durations. Using meta-analytic approaches, we show that, on average, dive durations decreased by 11% with every 1°C increase in water temperature. Larger increases in temperature (e.g. +3°C versus +8-9°C) exerted stronger effects on dive durations. Although species that respire bimodally are projected to be more resilient to the effects of temperature on dive durations than purely aerial breathers, we found no significant difference between these groups. Body mass had a weak impact on mean dive durations, with smaller divers being impacted by temperature more strongly. Few studies have examined thermal phenotypic plasticity ( N =4) in diving ectotherms, and all report limited plasticity. Average water temperatures in marine and freshwater habitats are projected to increase between 1.5 and 4°C in the next century, and our data suggest that this magnitude of warming could translate to substantial decreases in dive durations, by approximately 16-44%. Together, these data shed light on an overlooked threat to diving ectothermic vertebrates and suggest that time available for underwater activities, such as predator avoidance and foraging, may be shortened under future warming., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

From a terrestrial ancestry, the fin whale ( Balaenoptera physalus ) is one of the largest animals on Earth with a sprinter anti-predator strategy, and a characteristic feeding mode, lunge feeding, which involves bouts of high-intensity muscle activity demanding high metabolic output. We investigated the locomotor muscle morphology and metabolism of this cetacean to determine whether its muscle profile (1) explains this unique swimming performance and feeding behaviour, (2) is or is not homogeneous within the muscle, and (3) predicts allometric variations inherent to an extreme body size. A predominantly fast-glycolytic phenotype characterized the fin whale locomotor muscle, composed of abundant fast-twitch (type IIA) fibres with high glycolytic potential, low oxidative capacity, relatively small size, and reduced number of capillaries. Compared with superficial areas, deep regions of this muscle exhibited a slower and more oxidative profile, suggesting a division of labour between muscle strata. As expected, the fin whale locomotor muscle only expressed the two slowest myosin heavy chain isoforms (I and IIA). However, it displayed anaerobic (glycolytic) and aerobic (lipid-based metabolism) capabilities higher than would be predicted from the allometric perspective of its extreme body size. Relationships between muscle metabolism and body mass were fibre-type specific. The 'sprinter' profile of the fin whale swimming muscle, particularly of its superficial compartment, supports physiological demands during both high-speed swimming and the lunge, when energy expenditure reaches maximal or supramaximal levels. Comparatively, the slower and more oxidative profile of the deep compartment of this muscle seems to be well designed for sustained, low-intensity muscle activity during routine swimming., Competing Interests: Competing interestsThe author declares no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)