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Metabolic consequences of submersion asphyxia in mammals and birds.

On submersion, the cardiovascular system of naturally diving animals is virtually transformed into a heart-brain-lung preparation as the result of intense and highly selective neurogenic vasoconstriction. A supply of oxygen from the circulating blood cells to the heart and the brain is thereby secured. Peripheral tissues, e.g. muscles and kidneys, have to depend on local stores of oxygen, i.e. myoglobin, or, when these are exhausted, on anaerobic metabolism. In spite of such physiological adjustments, however, arterial pO2 will inevitably decrease throughout the underwater episode. In prolonged dives values lower than 10mmHg can be observed. Moreover, the ischaemia in the peripheral tissues will, in some animals, e.g. arctic seals and whales, result in a profound decrease in the temperature of tissues such as the skin. The aim of the present report is to review some of the more important biochemical mechanisms that allow diving mammals and birds to cope with the above-mentioned combined asphyxic and temperature stresses for prolonged periods. Some of the osmotic problems arising from the rapid wash-out of acidic metabolic end products from the previously ischaemic tissues on emergence are also discussed.

Adaptation, Physiological↗

Decompression from a deep nitrogen/oxygen saturation dive--a case report.

Ten divers participated in a 4.5 d nitrogen/oxygen saturation dive to 165 fsw. There were daily 2 h excursions to 61 msw (200 fsw). The divers breathed air during the excursions and 0.51 bar (0.5 atm) oxygen in nitrogen at 50.3 msw (165 fsw). The final decompression began 6 h after the last excursion. The oxygen partial pressure was 0.51 bar (0.5 atm) from 50.3 to 13.7 msw (165 to 45 fsw), and air was used from 13.7 msw (45 fsw) to the surface. By 6.1 msw (20 fsw), four divers had developed decompression sickness. A fifth diver developed decompression sickness during a commercial air flight 68 h after surfacing. Comparison of ascent rates for this dive and for air or nitrogen/oxygen saturation dives reported in the literature suggests that deeper dives require slower rates of ascent. Dives shallower than 30.5 msw (100 fsw) had a mean ascent rate of 1 msw/h (3.2 fsw/h) and 14 decompression incidents in 107 man-exposures. Dives deeper than 30.5 msw (100 fsw) had a mean rate of 0.76 msw/h (2.5 fsw/h) and 14 incidents in 45 man-exposures.

Adaptation, Physiological↗

Physiological responses to exercise at 47 and 66 ATA.

Five male volunteers served as subjects for exercise studies during three dives to pressures of 47 and 66 ATA while breathing gases containing 0.5 ATA PO2 and varying amounts of N2 and He. The inspired gas density ranged from 1.1 g/l (BTPS) at the surface to 17.1 g/l at the highest pressure. Dyspnea at rest and during exercise was evident in all divers and was predominantly inspiratory in nature. Despite the dyspnea, divers were able to perform work requiring an O2 consumption larger than 2 l/min STPD at each depth. Compared with surface measurements, moderate work at depth was associated with alveolar hypoventilation, arterial hypercapnia, very large physiological dead space, and higher levels of arterial lactate and signs of simultaneous respiratory and metabolic acidosis. The increase of ventilation that accompanies the onset of acidemia at the surface was not present at depth. Acidemia at depth was more severe, and its onset occurred at lesser work rates than at 1 ATA. No large differences could be ascertained when a variety of responses obtained with inspired gas having a density of 7.9 g/l at 47 ATA were compared with those obtained with an inspired gas density of 17.1 g/l at 66 ATA. It appears that the major impact of the environment on the physiological responses to work was almost fully manifested at a pressure of 47 ATA with a He-O2 gas mixture. It is cautioned that maximum work tolerance may be an insufficient assessment of the physiological condition of a diver exposed to these high pressures.

Acid-Base Equilibrium↗

Foraging energetics and diving behavior of lactating New Zealand sea lions, Phocarctos hookeri.

The New Zealand sea lion, Phocarctos hookeri, is the deepest- and longest-diving sea lion. We were interested in whether the diving ability of this animal was related to changes in its at-sea and diving metabolic rates. We measured the metabolic rate, water turnover and diving behavior of 12 lactating New Zealand sea lions at Sandy Bay, Enderby Island, Auckland Islands Group, New Zealand (50 degrees 30'S, 166 degrees 17'E), during January and February 1997 when their pups were between 1 and 2 months old. Metabolic rate (rate of CO(2) production) and water turnover were measured using the (18)O doubly-labeled water technique, and diving behavior was measured with time/depth recorders (TDRs). Mean total body water was 66.0+/-1.1 % (mean +/- s.d.) and mean rate of CO(2) production was 0. 835+/-0.114 ml g(-)(1 )h(-)(1), which provides an estimated mass-specific field metabolic rate (FMR) of 5.47+/-0.75 W kg(-)(1). After correction for time on shore, the at-sea FMR was estimated to be 6.65+/-1.09 W kg(-)(1), a value 5.8 times the predicted standard metabolic rate of a terrestrial animal of equal size. The mean maximum dive depth was 353+/-164 m, with a mean diving depth of 124+/-36 m. The mean maximum dive duration was 8.3+/-1.7 min, with an average duration of 3.4+/-0.6 min. The deepest, 550 m, and longest, 11.5 min, dives were made by the largest animal (155 kg). Our results indicate that the deep and long-duration diving ability of New Zealand sea lions is not due to a decreased diving metabolic rate. Individual sea lions that performed deeper dives had lower FMRs, which may result from the use of energetically efficient burst-and-glide locomotion. There are differences in the foraging patterns of deep and shallow divers that may reflect differences in surface swimming, time spent on the surface and/or diet. Our data indicate that, although New Zealand sea lions have increased their O(2) storage capacity, they do not, or cannot, significantly reduce their at-sea metabolic rates and are therefore likely to be operating near their physiological maximum.

Animals↗

History of polar whaling: insights into the physiology of the great whales.

The sheer size and pelagic nature of the great whales has effectively precluded detailed studies of most of their physiological processes. The vast majority of all data for these species have come from anatomical studies conducted on specimens that were caught in commercial and native whaling operations. In both the polar regions, an incredible number of whales were hunted, but anatomical studies were not usually conducted until relatively recent times. However, the anatomical data that do exist provide a valuable insight into some of the physiological demands placed on the animals by their marine habitat. These include information on blubber and nutrition; baleen and feeding ecology; contaminant chemistry and tissue samples; diving chemistry and acoustics. Taken together, these anatomical data provide the only substantial information on how these animals dive and hunt. Recent breakthroughs in chemical techniques however, are providing even greater details on function (for example, fatty acid signature methods). Coupled with advanced methods for tracking these whales at sea (acoustic and satellite), future studies should provide significant new information on the general physiology of these difficult to study species.

Adipose Tissue↗

Evolution of human hypoxia tolerance physiology.

Analysis of human responses to hypobaric hypoxia in different lineages (lowlanders, Andean natives, Himalayan natives, and East Africans) indicates 'conservative' and 'adaptable' physiological characters involved in human responses to hypoxia. Conservative characters, arising by common descent, dominant and indeed define human physiology, but in five hypoxia response systems analyzed, we also found evidence for 'adaptable' characters at all levels of organization in all three high altitude lineages. Since Andeans and Himalayans have not shared common ancestry with East Africans for most of our species history, we suggest that their similar hypoxia physiology may represent the 'ancestral' condition for humans--an interpretation consistent with recent evidence indicating that our species evolved under 'colder, drier, and higher' conditions in East Africa where the phenotype would be simultaneously advantageous for endurance performance and for high altitude hypoxia. It is presumed that the phenotype was retained in low capacity form in highlanders and in higher capacity form in most lowland lineages (where it would be recognized by most physiologists as an endurance performance phenotype). Interestingly, it is easier for modern molecular evolution theory to account for the origin of 'adaptable' characters through positive selection than for conserved traits. Many conserved physiological systems are composed of so many gene products that it seems difficult to account for their unchanging state (for unchanging structure and function of hundreds of proteins linked in sequence to form the physiological system) by simple models of stabilizing selection.

Acclimatization↗

Effects of underwater sound exposure on neurological function and brain histology.

To evaluate the safety of sonar exposure from a neurological perspective, the vulnerability of the central nervous system to underwater exposure with high-intensity, low-frequency sound (HI-LFS) was experimentally examined. Physiological, behavioral and histological parameters were measured in anesthetized, ventilated rats exposed to brief (5 min), underwater HI-LFS. Exposure to 180 dB sound pressure level (SPL) re 1 microPa at 150 Hz (n = 9) did not alter acute cardiovascular physiology (arterial blood pH, pO(2), pCO(2), heart rate, or mean arterial blood pressure) from that found in controls (n = 11). Rats exposed to either 180 dB SPL re 1 microPa at 150 Hz (n = 12) or 194 dB SPL re 1 microPa at 250 Hz (n = 12) exhibited normal cognitive function at 8 and 9 days after sound exposure. Evaluation of neurological motor function revealed a minor deficit 7 days after 180 dB SPL/150 Hz exposure that resolved by 14 days, and no deficits after 194 dB SPL/250 Hz exposure. No overt histological damage was detected in any group. These data suggest that underwater HI-LFS exposure may cause transient, mild motor dysfunction.

Analysis of Variance↗

Increased pulmonary vascular resistance and reduced stroke volume in association with CO2 retention and inferior vena cava dilatation.

Changes in cardiovascular parameters elicited during a maximal breath hold are well described. However, the impact of consecutive maximal breath holds on central hemodynamics in the postapneic period is unknown. Eight trained apnea divers and eight control subjects performed five successive maximal apneas, separated by a 2-min resting interval, with face immersion in cold water. Ultrasound examinations of inferior vena cava (IVC) and the heart were carried out at times 0, 10, 20, 40, and 60 min after the last apnea. The arterial oxygen saturation level and blood pressure, heart rate, and transcutaneous partial pressures of CO(2) and O(2) were monitored continuously. At 20 min after breath holds, IVC diameter increased (27.6 and 16.8% for apnea divers and controls, respectively). Subsequently, pulmonary vascular resistance increased and cardiac output decreased both in apnea divers (62.8 and 21.4%, respectively) and the control group (74.6 and 17.8%, respectively). Cardiac output decrements were due to reductions in stroke volumes in the presence of reduced end-diastolic ventricular volumes. Transcutaneous partial pressure of CO(2) increased in all participants during breath holding, returned to baseline between apneas, but remained slightly elevated during the postdive observation period (approximately 4.5%). Thus increased right ventricular afterload and decreased cardiac output were associated with CO(2) retention and signs of peripheralization of blood volume. These results indicate that repeated apneas may cause prolonged hemodynamic changes after resumption of normal breathing, which may suggest what happens in sleep apnea syndrome.

Adaptation, Physiological↗

A dynamic study of vanilmandelic acid elimination in the urine of divers in a dive at 21 ATA in saturation with a helium-oxygen respiratory mixture.

The purpose of this paper is a first assessment of the effect of stress agents specific for hyperbarism in professional divers. We selected the elimination of vanilmandelic acid (VMA) in the urine as an indicator. We studied four professional divers in a dive at 21 ATA. In order to assess the strain-induced by every distinct moment of the saturation profile we took samples of urine and determined VMA at four-hour intervals, before and after the distinct moment of saturation. The values found in the same four divers at the surface were used as "controls". VMA dosing in the urine proves to be a sensitive indicator for the assessment of stress and adaptive reactions during saturation.

Administration, Inhalation↗