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Metabolic rates of captive grey seals during voluntary diving.

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.

Animals↗

Stroke volume and cardiac output in juvenile elephant seals during forced dives.

The aim of this study was to examine the effect of forced diving on cardiac dynamics in a diving mammal by evaluating cardiac output and heart rate. We used MR Imaging and phase contrast flow analysis to obtain accurate flow measurements from the base of the aorta. Heart rate (fh) and cardiac output (Q) were measured before, during and after dives in four restrained juvenile northern elephant seals, Mirounga angustirostris, and stroke volume (Vs) was calculated (Vs=Q/fh). Mean Q during diving (4011+/-387 ml min(-1)) and resting (6530+/-1018 ml min(-1)) was not significantly different (paired t-test; P<0.055). Diving was accompanied by a 20% increase in Vs over the pre-dive level. Pre-dive, post-dive or diving fh was not significantly correlated with Vs during any state. Diving Vs correlated negatively with the bradycardic ratio (diving fh to pre-dive fh). In this study, the degree of bradycardia during diving was less than in previous pinniped studies, suggesting that the reduction in vagal input may contribute to the observed increase in Vs.

Animals↗

Energetic costs of diving and thermal status in European shags (Phalacrocorax aristotelis).

Diving is believed to be very costly in cormorants (Phalacrocoracidae) when compared with other avian divers because of their poor insulation and less-efficient foot propulsion. It was therefore suggested that cormorants might employ a behavioural strategy to reduce daily energy expenditure by minimizing the amount of time spent in water. However, European shags (Phalacrocorax aristotelis) have been observed to spend up to 7 h day(-1) diving in water of around 5-6 degrees C. To gain a better understanding of the energetic requirements in European shags, we measured their metabolic rates when resting in air/water and during shallow diving using respirometry. To investigate the effects of water temperature and feeding status on metabolic rate, birds dived at water temperatures ranging from 5 to 13 degrees C in both post-absorptive and absorptive states. In parallel with respirometry, stomach temperature loggers were deployed to monitor body temperature. Basal metabolic rate (BMR) was almost identical to allometric predictions at 4.73 W kg(-1). Metabolic rate when resting on water, during diving and after feeding was significantly elevated when compared with the resting-in-air rate. During diving, the metabolic rate of post-absorptive shags increased to 22.66 W kg(-1), which corresponds to 4.8x BMR. Minimum cost of transport (COT) was calculated at 17.8 J kg(-1) m(-1) at a swim speed of 1.3 m s(-1). Feeding before diving elevated diving metabolic rate by 13% for up to 5 h. There was a significant relationship between diving metabolic rate and water temperature, where metabolic rate increased as water temperature declined. Thermal conductance when resting in air at 10-19 degrees C was 2.05 W m(-2) degrees C(-1) and quadrupled during diving (7.88 W m(-2) degrees C(-1)). Stomach temperature when resting in air during the day was 40.6 degrees C and increased during activity. In dive trials lasting up to 50 min, stomach temperature fluctuated around a peak value of 42.0 degrees C. Hence, there is no evidence that European shags might employ a strategy of regional hypothermia. The energetic costs during shallow diving in European shags are considerably lower than has previously been reported for great cormorants (Phalacrocorax carbo) and are comparable to other foot-propelled divers. The lower dive costs in shags might be the consequence of a more streamlined body shape reducing hydrodynamic costs as well as a greater insulative plumage air layer (estimated to be 2.71 mm), which reduces thermoregulatory costs. The latter might be of great importance for shags especially during winter when they spend extended periods foraging in cold water.

Animals↗

Effect of pharmacological blockade on cardiovascular responses to voluntary and forced diving in muskrats.

Neural control of free and forced diving bradycardia and peripheral resistance was studied in the muskrat (Ondatra zibethicus) by means of acute pharmacological blockade with the muscarinic blocker atropine, the alpha-adrenergic blocker phentolamine and the beta-adrenergic blockers nadolol and propranolol. Saline injection was used as a control. Heart rate in control animals increased before voluntary dives and dropped markedly as soon as the animals submerged. Heart rate started increasing towards the end of voluntary dives and reached pre-dive values within the first 5 s of recovery. Pre-dive and post-dive tachycardia were reduced in beta-blocked animals, emphasizing the role of the sympathetic system during the preparatory and recovery periods of voluntary dives. Diving bradycardia and the acceleration in heart rate before surfacing were abolished by atropine and unaffected by nadolol, demonstrating the importance of vagal efferent activity during diving. The results after blockade with nadolol suggest that there is an accentuated antagonism between the two branches of the autonomic nervous system during diving, so that parasympathetic influences on the heart predominate. Propranolol-treated muskrats had a higher diving heart rate than saline- and nadolol-treated animals, which may be due to a sedative effect caused by propranolol crossing the blood-brain barrier, a blockade of central catecholaminergic pathways or a peripheral neural effect, due to the anaesthetic properties of propranolol. Phentolamine did not affect diving bradycardia, indicating that diving bradycardia occurs independently of peripheral vasoconstriction.(ABSTRACT TRUNCATED AT 250 WORDS)

Adrenergic alpha-Antagonists↗

Venous bubble count declines during strenuous exercise after an open sea dive to 30 m.

INTRODUCTION: The effect of post-dive exercise on bubble formation remains controversial, although the current practice of divers and aviators is to avoid strenuous exercise after diving. Previously, we have shown that exercising 24 h before a dive, or during a decompression stop, significantly reduces bubble formation in man. The objective of this study was to determine whether a short period of strenuous post-dive exercise promotes venous bubble formation. METHODS: Seven male military divers performed an open-sea field dive to a maximum depth of 30 m for 30 min. At maximum depth, subjects performed mild underwater fin swimming, followed by standard decompression. Diving was followed by a post-dive exercise session consisting of short, strenuous incremental upright cycle ergometry, up to 85% of maximal oxygen uptake, for about 10 min. Subjects were monitored for venous gas bubbles in the right heart with an echo-imaging system starting 20 min post-dive while in the supine position, during cycle ergometry in the seated upright position, and immediately after exercise in a supine position. RESULTS: The average number of bubbles was 1.5 +/- 1.4 bubbles x cm(-2) 20 min after diving. Changes in posture from supine to seated upright resulted in significant reduction of bubbles to 0.6 +/- 1.3 bubbles x cm(-2) (p = 0.043), with further reduction to 0.2 +/- 0.3 bubbles x cm(-2) at the end of exercise (p = 0.02). No cases of DCS or intra-pulmonary shunt were observed during or following post-dive exercise. DISCUSSION: These results suggest that post-dive strenuous exercise after a single field dive reduces post-dive gas bubble formation in well-trained military divers. Additional findings are needed for normal sports divers.

Adult↗

Daily diving pattern of Korean and Japanese breath-hold divers (ama).

Daily diving patterns and thoracic skin and sea-water temperatures were recorded during the entire work shift of Korean female unassisted (cachido) and Japanese male unassisted and partly assisted (funado) divers using Underwater Physiological Data Loggers developed in Buffalo and Japan. All 3 groups of divers were studied during the summer of 1989 and 1990. Additional studies were conducted during the winter of 1991 on Korean female divers who, unlike Japanese divers, dive all year round. The water temperature of the diving grounds in summer was 24 degrees C in both Korea and Japan, and 10 degrees C during winter in Korea. Both Korean female and Japanese male cachido divers made 113-138 dives a day and stayed in the water a total of 170-200 min.day-1, of which only 52-63 min were spent diving submerged, and the remaining time at the water surface. These diving patterns were not different between female and male cachido divers. Compared with Japanese male divers, Korean female divers dived to a shallower depth (3.7 vs. 6.9 m) with shorter dive time (29 vs. 37 s) and shorter bottom time (14 vs. 18 s). Velocities of descent (0.72 vs. 0.47 m.s-1) and ascent (0.77 vs. 0.56 m.s-1) were also slower in female divers than in male divers. The diving pattern of Korean female divers was similar in both summer and winter. Although all cachido divers wore wet suits and thus were protected from severe cold stress, thoracic skin temperature decreased during a work shift by 7 degrees C in winter (vs. 1 degree C in summer) in Korean divers. Compared with Japanese male cachido divers, Japanese male funado divers stayed in the diving ground (including time in the boat) longer (201 vs. 305 min.day-1) but performed only 23 dives per day. The average diving depth (9.7 m), duration (69 s), and bottom time for each dive (45 s), however, were significantly greater in funados. The velocity of vertical descent (1.0 m.s-1) was also significantly greater in funados because they descend with a weight (8-12 kg). The rate of ascent was not different.

Body Temperature↗

Heart rates and diving behavior of leatherback sea turtles in the eastern pacific ocean

Heart rates and diving behavior of leatherback sea turtles (Dermochelys coriacea) were monitored at sea during the internesting interval. Instruments that recorded the electrocardiogram and the depth and duration of dives were deployed on six female leatherback turtles as they laid eggs at Playa Grande, Costa Rica. Turtles dived continually for the majority of the internesting interval and spent 57-68 % of the time at sea submerged. Mean dive depth was 19+/-1 m (mean +/- s.d.) and the mean dive duration was 7.4+/-0.6 min. Heart rate declined immediately upon submergence and continued to fall during descent. All turtles showed an increase in heart rate before surfacing. The mean heart rate during dives of 17.4+/-0.9 beats min-1 (mean +/- s.d.) was significantly lower than the mean heart rate at the surface of 24.9+/-1.3 beats min-1 (P<0.05). Instantaneous heart rates as low as 1.05 beats min-1 were recorded during a 34 min dive. The mean heart rate over the entire dive cycle (dive + succeeding surface interval; 19.4+/-1.3 beats min-1) was more similar to the heart rate during diving than to the heart rate at the surface. Although dive and surface heart rates were significantly different from each other, heart rates during diving were 70 % of heart rates at the surface, showing that leatherback turtles do not experience a dramatic bradycardia during routine diving.

Journal Article↗

Diving and pregnancy.

Scuba diving during pregnancy has increased in incidence as a result of substantial growth in the number of young females attracted to sport diving. This review summarizes the physiological changes induced by immersion, diving and decompression, on male and female divers. Furthermore, it extends to literature review, in animal models, of the susceptibility of a pregnant animal to diving decompression injury. Publications regarding reports of diving injury in pregnant humans are also reviewed, comprising very recent material from the sport diving community. It is concluded that there is no countraindication to diving for the normal, healthy, nonpregnant female. However, pregnant females should refrain from diving, because the fetus is not protected from decompression problems and is at risk of malformation and gas embolism after decompression disease. It is prudent to advise pregnant patients of the increased risk of diving problems for the fetus during pregnancy. However, should a woman have completed a dive during early pregnancy because she was unaware she was pregnant, the present evidence is not to recommend an abortion, because several normal pregnancies have been documented even if diving is continued. Snorkeling can still be practiced during pregnancy, but scuba diving should be discontinued until after the birth period.

Animals↗

Long term retention of safe diving skills.

This short report describes a 20-month follow-up of safe diving skills, extending the 8-month retention period previously published in this journal. Thirty-four recreational swimmers with poor diving skills were evaluated before and immediately after a diving skills intervention program. Twenty-two returned for the eight-month follow-up evaluation and 16 returned 20 months post. As with the earlier study, Treadwater, Deck, Block and Running dives were video-recorded, and maximum depth, distance, velocity, entry angle and flight distance were compared. Underwater hand and arm positions were examined. Pre-intervention, a breaststroke arm action before maximum depth occurred in 18% of all dives and 38% of Treadwater dives. This was eliminated post-intervention, improving head protection. The Treadwater dive elicited the greatest mean maximum depth, and ANOVA showed depth for this entry decreased (improved) following intervention and remained shallower at the eight-month and 20-month post follow-ups. The Block dive also became shallower following intervention while the Deck dive remained unchanged. As seven 10-minute skills sessions resulted in shallower dives with safer hand and arm positions, and these skills were retained over a 600 day non-practice period, it is reliable to consider that the inclusion of safe diving skills in learn-to-swim programs can provide a diving spinal cord injury prevention strategy.

Adult↗

Norwegian deep diving trials.

In 1983 NUTEC, together with two diving companies, completed two dives with 12 divers (6 in each dive) to pressures equivalent to 350 m s.w., one dive lasted for 17 d, and the other, 24 d. The purpose of the dives was to demonstrate that the diving companies were prepared for diving to 300 m depth in the North Sea. No major medical or physiological problems arose during the dives, although all divers had minor symptoms of high pressure nervous syndrome during compressions. During decompression three decompression sickness incidents occurred, which involved pain only, and all were successfully treated. All divers went through comprehensive medical physiological examinations before and after the dives. No significant changes from values measured before diving have been found in the six divers who have so far been examined after diving, except that five of them were considerably more sensitive to CO2 after the dive than before. Several problems arose in connection with the divers' breathing equipment, thermal protection and communication, which need to be improved.

Atmosphere Exposure Chambers↗

Ontogeny of diving behaviour in the Australian sea lion: trials of adolescence in a late bloomer.

1. Foraging behaviours of the Australian sea lion (Neophoca cinerea) reflect an animal working hard to exploit benthic habitats. Lactating females demonstrate almost continuous diving, maximize bottom time, exhibit elevated field metabolism and frequently exceed their calculated aerobic dive limit. Given that larger animals have disproportionately greater diving capabilities, we wanted to examine how pups and juveniles forage successfully. 2. Time/depth recorders were deployed on pups, juveniles and adult females at Seal Bay Conservation Park, Kangaroo Island, South Australia. Ten different mother/pup pairs were equipped at three stages of development (6, 15 and 23 months) to record the diving behaviours of 51 (nine instruments failed) animals. 3. Dive depth and duration increased with age. However, development was slow. At 6 months, pups demonstrated minimal diving activity and the mean depth for 23-month-old juveniles was only 44 +/- 4 m, or 62% of adult mean depth. 4. Although pups and juveniles did not reach adult depths or durations, dive records for young sea lions indicate benthic diving with mean bottom times (2.0 +/- 0.2 min) similar to those of females (2.1 +/- 0.2 min). This was accomplished by spending higher proportions of each dive and total time at sea on or near the bottom than adults. Immature sea lions also spent a higher percentage of time at sea diving. 5. Juveniles may have to work harder because they are weaned before reaching full diving capability. For benthic foragers, reduced diving ability limits available foraging habitat. Furthermore, as juveniles appear to operate close to their physiological maximum, they would have a difficult time increasing foraging effort in response to reductions in prey. Although benthic prey are less influenced by seasonal fluctuations and oceanographic perturbations than epipelagic prey, demersal fishery trawls may impact juvenile survival by disrupting habitat and removing larger size classes of prey. These issues may be an important factor as to why the Australian sea lion population is currently at risk.

Aging↗

Human sympathetic nerve activity to glabrous skin does not increase during simulated diving.

In humans, cardiovascular adjustment to simulated diving causes a marked increase in sympathetic outflow to intramuscular vessels and muscle vasoconstriction. Skin vasoconstriction in the hand also occurs during diving in humans. Skin nerve sympathetic activity (SSA), containing vasoconstrictor signals to glabrous skin, unexpectedly was reduced during diving in a previous study of SSA recorded in the peroneal nerve. SSA was recorded by microneurography in the median nerve in 13 healthy volunteers during simulated diving. Skin blood flow in the hand and one finger was monitored. The typical SSA response, irrespective of duration of diving and water temperature, was an increase during the control period immediately prior to immersion of the face and a sudden reduction of SSA when the face was immersed. The increase in SSA preceding the dive was accompanied by vasoconstriction, which continued during the dive, but re-dilation regularly occurred before the end of the dive. Inhibition of SSA was not total. Mental arithmetic during diving evoked strong bursts of SSA, similar to those seen normally during mental stress. It is concluded that the true response of SSA to simulated diving is an inhibition of the immediately preceding outflow, in agreement with observations of cutaneous blood flow in animals. The skin vasoconstriction recorded during simulated diving is a consequence of an SSA increase before the procedure, suggested to be a stress response before the forthcoming manoeuvre. The SSA response during simulated diving is the opposite to that of sympathetic outflow to muscle, which emphasizes the diversity of sympathetic regulation of different organ systems.

Adult↗

Hemoglobin concentrations and blood gas tensions of free-diving Weddell seals.

Arterial blood gas tensions, pH, and hemoglobin concentrations were measured in four free-diving Weddell seals Leptonychotes weddelli. A microprocessor-controlled sampling system enabled us to obtain 24 single and 31 serial aortic blood samples. The arterial O2 tension (PaO2) at rest [78 +/- 13 (SD) Torr] increased with diving compression to a maximum measured value of 232 Torr and then rapidly decreased to 25-35 Torr. The lowest diving PaO2 we measured was 18 Torr just before the seal surfaced from a 27-min dive. A consistent increase of arterial hemoglobin concentrations from 15.1 +/- 1.10 to 22.4 +/- 1.41 g/100 ml (dives less than 17 min) and to 25.4 +/- 0.79 g/100 ml (dives greater than 17 min) occurred during each dive. We suggest that an extension of the sympathetic outflow of the diving reflex possibly caused profound contraction of the Weddell seal's very large spleen (0.89% of body wt at autopsy), although we have no direct evidence. This contraction may have injected large quantities of red blood cells (2/3 of the total) into the seal's central circulation during diving and allowed arterial O2 content to remain constant for the first 15-18 min of long dives. The increase of arterial CO2 tensions during the dive and the compression increase of arterial N2 tensions were also moderated by injecting red blood cells sequestered at ambient pressure. After each dive circulating red blood cells are oxygenated and rapidly sequestered, possibly in the spleen during the first 15 min of recovery.

Animals↗

Surface pauses in relation to dive duration in imperial cormorants; how much time for a breather?

Air-breathing animals diving to forage can optimize time underwater by diving with just enough oxygen for the projected performance underwater. By so doing they surface with minimal body oxygen levels, which leads to maximal rates of oxygen uptake. We examined whether imperial cormorants Phalacrocorax atriceps adhere to this by examining dive:pause ratios in birds diving for extended, continuous periods to constant depths, assuming that the oxygen used underwater was exactly replenished by the periods at the surface. Examination of the cumulative time spent in surface pauses relative to the cumulative time spent in diving showed that surface pauses increase according to a power curve function of time spent in the dive or water depth. In a simplistic model we considered the rate at which birds expended energy underwater to be constant and that the rate of oxygen replenishment during the surface pause was directly proportional to the oxygen deficit. We then worked out values for the rate constant for the surface pause before using this constant to examine bird body oxygen levels immediately pre- and post dive. The model predicted that imperial cormorants do not submerge with just enough oxygen to cover their projected dive performance but rather dive with substantial reserves, although these reserves decrease with increasing dive depth/duration. We speculate that these oxygen reserves may be used to enhance bird survival when rare events, such as the appearance of predators or discovery of large prey requiring extended handling time, occur. The form of the oxygen saturation curve over time at the surface means that the time costs for maintaining constant oxygen reserves become particularly onerous for long, deep dives, so the observed decrease in reserves with increasing dive duration is expected in animals benefiting by optimizing for time.

Animals↗

Regulation of stroke pattern and swim speed across a range of current velocities: diving by common eiders wintering in polynyas in the Canadian Arctic.

Swim speed during diving has important energetic consequences. Not only do costs increase as drag rises non-linearly with increasing speed, but speed also affects travel time to foraging patches and therefore time and energy budgets over the entire dive cycle. However, diving behaviour has rarely been considered in relation to current velocity. Strong tidal currents around the Belcher Islands, Nunavut, Canada, produce polynyas, persistent areas of open water in the sea ice which are important habitats for wildlife wintering in Hudson Bay. Some populations of common eiders Somateria mollissima sedentaria remain in polynyas through the winter where they dive to forage on benthic invertebrates. Strong tidal currents keep polynyas from freezing, but current velocity can exceed 1.5 m s(-1) and could influence time and energy costs of diving and foraging. Polynyas therefore provide naturally occurring flume tanks allowing investigation of diving strategies of free ranging birds in relation to current velocity. We used a custom designed sub-sea ice camera to non-invasively investigate over 150 dives to a depth of 11.3 m by a population of approximately 100 common eiders at Ulutsatuq polynya during February and March of 2002 and 2003. Current speed during recorded dives ranged from 0 to 1 m s(-1). As currents increased, vertical descent speed of eiders decreased, while descent duration and the number of wing strokes and foot strokes during descent to the bottom increased. However, nearly simultaneous strokes of wings and feet, and swim speed relative to the moving water, were maintained within a narrow range (2.28+/-0.23 Hz; 1.25+/-0.14 m s(-1), respectively). This close regulation of swim speed over a range in current speed of 1.0 m s(-1) might correspond to efficient muscle contraction rates, and probably reduces work rates by avoiding rapidly increasing drag at greater speeds; however, it also increases travel time to benthic foraging patches. Despite regulation of average swim speed, high instantaneous speeds during oscillatory stroking can increase dive costs due to drag. While most diving birds have been considered either foot or wing propelled, eider ducks used both wing and foot propulsion during descent. Our observations indicate that the power phase of foot strokes coincides with the transition between upstroke and downstroke of the wings, when drag is greatest. Coordinated timing between foot and wing propulsion could therefore serve to maintain a steadier speed during descent and decrease the costs of diving. Despite tight regulation of stroke and swim speed patterns, descent duration and total number of foot and wing strokes during descent increase non-linearly with increasing current velocity, suggesting an increase in energetic costs of diving.

Animals↗

Extreme diving of beaked whales.

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 min; Md: 47 min) 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 min for Zc and 92 min 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.

Activity Cycles↗

Diving energetics in king penguins (Aptenodytes patagonicus).

Dive duration in wild king penguins and the energetic cost of swimming in a 30m long swim channel were determined at Ile de la Possession, Crozet Archipelago, using external data loggers and respirometry, respectively. Calibrated electronic data loggers equipped with a pressure sensor were used to determine dive durations: 95% of dives were less than 6 min long and 66% of dives were less than 4 min long. Dive patterns show that king penguins may intersperse long dive durations (4-6.3 min) with short ones (1.5-3 min) and make surface pauses of variable duration between them (0.5-3.5 min), or dive regularly (for up to 5 h) with long dive durations (5 min) and constant interdive surface intervals (1.5 min). The latter indicates that the aerobic dive limits (ADL) of this species could be higher and oxygen consumption lower than previously reported. Assuming that king penguins dive within their aerobic limit, different approaches to the analysis of the data obtained in the swim channel are discussed to derive the ADL. Swimming speeds observed in the channel ranged from 0.9 to 3.4 m s-1. Transport costs were lowest between 1.8 and 2.2 m s-1. Although at 2.2 m s-1 king penguins used only 10.3 Wkg-1 over a dive+surface cycle (minimal transport costs of 4.7 J kg-1 m-1), we speculate that tisse oxygen consumption during submergence may be as low as 0.23 ml O2 kg-1 s-1 (2.1 times standard metabolic rate, SMR) or perhaps lower (which gives an ADL of 4.2 min). During surface phases, oxygen uptake would be increased to at least 1 ml O2kg-1 s-1 (9.3 times SMR). This implies that at least 70% of all dives are aerobic. Potential physiological mechanisms allowing king penguins to partition O2 consumption between submergence and surface periods remain, however, unclear.

Animals↗

The diving physiology of bottlenose dolphins (Tursiops truncatus). II. Biomechanics and changes in buoyancy at depth.

During diving, marine mammals must balance the conservation of limited oxygen reserves with the metabolic costs of swimming exercise. As a result, energetically efficient modes of locomotion provide an advantage during periods of submergence and will presumably increase in importance as the animals perform progressively longer dives. To determine the effect of a limited oxygen supply on locomotor performance, we compared the kinematics and behavior of swimming and diving bottlenose dolphins. Adult bottlenose dolphins (Tursiops truncatus) were trained to swim horizontally near the water surface or submerged at 5 m and to dive to depths ranging from 12 to 112 m. Swimming kinematics (preferred swimming mode, stroke frequency and duration of glides) were monitored using submersible video cameras (Sony Hi-8) held by SCUBA divers or attached to a pack on the dorsal fin of the animal. Drag and buoyant forces were calculated from patterns of deceleration for horizontally swimming and vertically diving animals. The results showed that dolphins used a variety of swimming gaits that correlated with acceleration. The percentage of time spent gliding during the descent phase of dives increased with depth. Glide distances ranged from 7.1+/-1.9 m for 16 m dives to 43.6+/-7.0 m (means +/- s.e.m.) for 100 m dives. These gliding patterns were attributed to changes in buoyancy associated with lung compression at depth. By incorporating prolonged glide periods, the bottlenose dolphin realized a theoretical 10-21 % energetic savings in the cost of a 100 m dive in comparison with dives based on neutral buoyancy models. Thus, modifying locomotor patterns to account for physical changes with depth appears to be one mechanism that enables diving mammals with limited oxygen stores to extend the duration of a dive.

Animals↗