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Myocardial blood flow and metabolism in the diving seal.

The adaptations of myocardial metabolism to diving asphyxia have been studied in 12 harbor seals (Phoca vitulina). Unanesthetized animals were submerged for periods of 10-16 min. Heart rate decreased from 135 to 12 beats/min. Myocardial blood flow decreased to an average of 10% of predive values and remained constant during the dive. The progressive reduction in arterial O2 content was associated with an increase in myocardial lactate and hydrogen ion production, but no change in glucose or free fatty acid extraction occurred. After restoration of breathing a reactive myocardial hyperemia and an immediate return to myocardial uptake of lactate were observed. Despite increased glycogenolytic activity throughout the dive, coronary flow distribution was fully controlled, and no evidence of ischemic dilatation of the left ventricle or S-T segment elevation in the electrocardiogram was observed. These adaptations to diving asphyxia in the seal myocardium permit a reduction of coronary blood flow comparable to that observed in the infarcted dog myocardium and therefore have relevance for therapeutic approaches to reduction of myocardial ischemic injury in humans.

Animals↗

Metabolic responses to forced dives in Pekin duck measured by indirect calorimetry and 31P-MRS.

We tested the hypothesis that forced-dived ducks experience a reduction in metabolic rate during prolonged submergence. Unidirectionally ventilated conscious ducks were subjected to forced dives by temporarily stopping the airflow in the ventilation system and simultaneously filling a face mask with cold water. A typical cardiovascular response to submergence was observed: bradycardia and maintained arterial blood pressure. Phosphorylated metabolite concentrations in the pectoral muscle were measured noninvasively by phosphorus magnetic resonance spectroscopy (31P-MRS). ATP content was constant, and phosphocreatine was depleted via the creatine kinase reaction at a rate similar to the resting rate of ATP turnover, which was estimated to be 0.9 mumol.min-1 x g-1 in resting perfused pectoral muscle of pentobarbital-anesthetized ducks. Oxygen from myoglobin supplied at most 12% of the ATP required by the resting muscle during dives. Whole animal postdive excess oxygen consumption and blood lactic acid accumulation suggested that the shortfall in aerobic metabolism during forced dives was compensated by an increase in anaerobic metabolism.

Adenosine Triphosphate↗

Temperature effect on the human dive response in relation to cold water near-drowning.

To facilitate analysis of mechanisms involved in cold water near-drowning, maximum breath-hold duration (BHD) and diving bradycardia were measured in 160 humans who were submerged in water temperatures from 0 to 35 degrees C at 5 degrees C intervals. For sudden submersion BHD was dependent on water temperature (Tw) according to the equation BHD = 15.01 + 0.92Tw. In cold water (0-15 degrees C), BHD was greatly reduced, being 25-50% of the presubmersion duration. BHD after brief habituation to water temperature and mild, voluntary hyperventilation was more than double that of sudden submersion and was also dependent on water temperature according to the equation BHD = 38.90 + 1.70Tw. Minimum heart rate during both types of submersions (diving bradycardia) was independent of water temperature. The results are pertinent to accidental submersion in cold water and show that decreased breath-holding capacity caused by peripheral cold stimulation reduces the effectiveness of the dive response and facilitates drowning. These findings do not support the postulate that the dive response has an important role in the enhanced resuscitatibility associated with cold water near-drowning, thereby shifting emphasis to hypothermia as the mechanism for this phenomenon.

Adult↗

Brain amino acid concentrations during diving and acid-base stress in turtles.

To assess the role of brain amino acid neurotransmitters in the breath hold of diving animals, concentrations of free amino acids present in the brains of turtles immediately after 2 h of apneic diving (at 20 degrees C) were measured. Additionally, the same measurements were performed on four other groups of animals subjected to 2 h of hypercapnia (8% CO2 in air), anoxia (N2 breathing), anoxia plus hypercapnia (8% CO2-92% N2), or air breathing (control). Significant changes in the concentrations of the inhibitory amino acid neurotransmitters known to affect respiration [gamma-aminobutyric acid (GABA) and taurine] were seen. GABA increased significantly in those animals subjected to anoxia, whereas taurine decreased significantly in the diving animals and increased significantly in those subjected to anoxia plus hypercapnia. These results suggest that the attenuated central ventilatory drive during diving in these animals may be related to alterations in brain concentrations of GABA and taurine.

Acid-Base Imbalance↗

Energetics of wet-suit diving in Japanese male breath-hold divers.

The present study was undertaken to investigate energy balance in professional male breath-hold divers in Tsushima Island, Japan. In 4 divers, rectal (Tre) and mean skin (Tsk) temperatures and rate of O2 consumption (VO2) were measured during diving work in summer (27 degrees C water) and winter (14 degrees C water). Thermal insulation and energy costs of diving work were estimated. In summer, comparisons were made of subjects clad either in wet suits (protected) or in swimming trunks (unprotected), and in winter, they wore wet suits. The average Tre in unprotected divers decreased to 36.4 +/- 0.2 degrees C at the end of 1-h diving work, but in protected divers it decreased to 37.2 +/- 0.3 degrees C in 2 h in summer and to 36.9 +/- 0.1 degree C in 1.5 h in winter. The average Tsk of unprotected divers decreased to 28.0 +/- 0.6 degrees C in summer and that of protected divers decreased to 32.9 +/- 0.5 degrees C in summer and 28.0 +/- 0.3 degrees C in winter. Average VO2 increased 190% (from 370 ml/min before diving to 1,070 ml/min) in unprotected divers in summer, but in protected divers it rose 120% (from 360 to 780 ml/min) in summer and 110% (from 330 to 690 ml/min) in winter. Overall thermal insulation (tissue and wet suit) calculated for protected divers was 0.065 +/- 0.006 degree C X kcal-1 X m-2 X h-1 in summer and 0.135 +/- 0.019 degree C X kcal-1 X m-2 X h-1 in winter.(ABSTRACT TRUNCATED AT 250 WORDS)

Acclimatization↗

Exercise tolerance and pulmonary gas exchange after deep saturation dives.

Pulmonary function and exercise tolerance were measured before and after three saturation dives to a pressure of 3.7 MPa. The atmospheres were heliox with partial pressures of oxygen of 40 kPa during the bottom phase and 50 kPa during the compression and decompression phase. The bottom times were 3, 10, and 13 days. Decompression time was 13 days. Precordial Doppler monitoring was done daily during the decompression, and an estimate of the total bubble load on the pulmonary circulation was calculated as the accumulated sum of bubble scores recorded for each diver. Nine of the 18 divers had chest symptoms with retrosternal discomfort or nonproductive cough after the dive. There were no changes in dynamic lung volumes. Transfer factor for carbon monoxide was significantly reduced from 12.3 +/- 1.2 to 10.9 +/- 1.3 mmol.kPa-1.min-1 (P less than 0.01), and maximum oxygen uptake was reduced from 3.98 +/- 0.36 to 3.42 +/- 0.37 l/min STPD (P less than 0.01) after the dives. Resting heart rate was increased from 64 +/- 6 to 75 +/- 8 min-1 (P less than 0.01). The ventilatory requirements in relation to oxygen uptake and carbon dioxide elimination were significantly increased (P less than 0.01) after the dives. The physiological dead space fraction of tidal volume was significantly higher and showed an increase with larger tidal volumes (P less than 0.05). Anaerobic threshold estimated from gas exchange data decreased from an oxygen uptake of 2.30 +/- 0.25 to 1.95 +/- 0.28 l/min STPD (P less than 0.05).(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

Alveolar gas composition and exchange during deep breath-hold diving and dry breath holds in elite divers.

End tidal O2 and CO2 (PETCO2) pressures, expired volume, blood lactate concentration ([Lab]), and arterial blood O2 saturation [dry breath holds (BHs) only] were assessed in three elite breath-hold divers (ED) before and after deep dives and BH and in nine control subjects (C; BH only). After the dives (depth 40-70 m, duration 88-151 s), end-tidal O2 pressure decreased from approximately 140 Torr to a minimum of 30.6 Torr, PETCO2 increased from approximately 25 Torr to a maximum of 47.0 Torr, and expired volume (BTPS) ranged from 1.32 to 2.86 liters. Pulmonary O2 exchange was 455-1,006 ml. CO2 output approached zero. [Lab] increased from approximately 1.2 mM to at most 6.46 mM. Estimated power output during dives was 513-929 ml O2/min, i.e. approximately 20-30% of maximal O2 consumption. During BH, alveolar PO2 decreased from approximately 130 to less than 30 Torr in ED and from 125 to 45 Torr in C. PETCO2 increased from approximately 30 to approximately 50 Torr in both ED and C. Contrary to C, pulmonary O2 exchange in ED was less than resting O2 consumption, whereas CO2 output approached zero in both groups. [Lab] was unchanged. Arterial blood O2 saturation decreased more in ED than in C. ED are characterized by increased anaerobic metabolism likely due to the existence of a diving reflex.

Adult↗

Psychophysiological reactions in humans during an open sea dive to 500 m with a hydrogen-helium-oxygen mixture.

Six commercial divers were investigated for neurological and psychosensorimotor responses during an open sea dive to 500 m with a hydrogen-helium-oxygen mixture containing 49% hydrogen. Results showed only moderate neurological symptoms of high-pressure nervous syndrome, whereas the narcotic effect of hydrogen was detectable, as investigated by psychosensorimotor tests. Nevertheless, the divers successfully carried out the main purpose of the operational dive, which was to prove the feasability of such diving methods by connecting specific elements of an offshore oil installation. Finally, these data support the hypothesis that hydrogen can alleviate some of the symptoms of the high-pressure nervous syndrome and can constitute a useful gas for commercial diving, as it decreases the density of the breathing mixture and therefore improves the living conditions, work, and comfort of the divers. Nevertheless, the present results underscore the relevance of research on individual susceptibility to pressure environment regardless of the composition of the breathing mixture.

Adult↗

Splenic contraction, catecholamine release, and blood volume redistribution during diving in the Weddell seal.

The spleen of the Weddell seal (Leptonychotes weddelli) may contract and inject red blood cells (RBCs) into the peripheral circulation during diving, but evidence for this hypothesis is indirect. Accordingly, we measured splenic dimensions by ultrasonography, plasma catecholamine concentrations, hemoglobin concentration, and hematocrit in five Weddell seals before and after intravenous epinephrine during halothane anesthesia and while awake at the surface after voluntary dives. Spleen size was reduced immediately after epinephrine injection or after the seal surfaced. Within the first 2 min after the seal surfaced, cephalocaudal splenic length was 71 +/- 2% (mean +/- SD; P < 0.05) and splenic thickness was 71 +/- 4% (P < 0.05) of the maximal resting values. Splenic size increased (half-time = 6-9 min) after the seal surfaced and was inversely correlated with plasma epinephrine and norepinephrine concentrations. Hemoglobin concentration increased from 17.5 +/- 5.3 g/dl (measured during general anesthesia) to 21.9 +/- 3.7 g/dl (measured in the first 2 min after surfacing). At these same times, the hematocrit increased from 44 +/- 12 to 55 +/- 8%. These values decreased (half-time = 12-16 min) after the seal surfaced. We estimate 20.1 liters of RBCs were sequestered at rest, presumably in the spleen, and released either on epinephrine injection or during diving. Catecholamine release and splenic contraction appear to be an integral part of the voluntary diving response of Weddell seals.

Animals↗

Decompression sickness in the rat following a dive on trimix: recompression therapy with oxygen vs. heliox and oxygen.

Trimix (a mixture of helium, nitrogen, and oxygen) has been used in deep diving to reduce the risk of high-pressure nervous syndrome during compression and the time required for decompression at the end of the dive. There is no specific recompression treatment for decompression sickness (DCS) resulting from trimix diving. Our purpose was to validate a rat model of DCS on decompression from a trimix dive and to compare recompression treatment with oxygen and heliox (helium-oxygen). Rats were exposed to trimix in a hyperbaric chamber and tested for DCS while walking in a rotating wheel. We first established the experimental model, and then studied the effect of hyperbaric treatment on DCS: either hyperbaric oxygen (HBO) (1 h, 280 kPa oxygen) or heliox-HBO (0.5 h, 405 kPa heliox 50%-50% followed by 0.5 h, 280 kPa oxygen). Exposure to trimix was conducted at 1,110 kPa for 30 min, with a decompression rate of 100 kPa/min. Death and most DCS symptoms occurred during the 30-min period of walking. In contrast to humans, no permanent disability was found in the rats. Rats with a body mass of 100-150 g suffered no DCS. The risk of DCS in rats weighing 200-350 g increased linearly with body mass. Twenty-four hours after decompression, death rate was 40% in the control animals and zero in those treated immediately with HBO. When treatment was delayed by 5 min, death rate was 25 and 20% with HBO and heliox, respectively.

Animals↗

Scuba diving with cochlear implants.

We report on a patient with bilateral cochlear implants (a Med-El Combi40 and a Med-El Combi40+), as well as considerable experience in scuba diving with both of his implants. After having been exposed to 68 and 89 dives, respectively, in depths of up to 43 m, both cochlear implants are in working order and the patient continues to receive excellent speech recognition scores with both cochlear implant systems. The presented data show that scuba diving after cochlear implantation is possible over a considerable number of dives without any major negative impact on the implants.

Cochlear Implants↗

Diving accidents in sports divers in Orkney waters.

Scapa Flow in Orkney is one of the major world centres for wreck diving. Because of the geography of Orkney and the nature of the diving, it is possible to make relatively accurate estimates of the number of dives taking place. The denominator of dive activity allows the unusual opportunity of precise calculation of accident rates. In 1999, one in every 178 sports divers visiting Orkney was involved in a significant accident, in 2000 the figure was one in 102. Some of these accidents appear to have been predictable and could be avoided by better education and preparation of visiting divers.

Athletic Injuries↗

Complications of saturation diving.

The experience of 458 man-dives with 731 excursions between 50 m and 300 m carried out by Royal Navy saturation divers is summarized. During saturation decompression there were 6 treated bends and 33 reported niggles. Two bends occurred in dives deeper than 249 m and the remaining 4 bends occurred in dives where decompression began in much less than the saturation stop time after completion of downward excursions. There was one case of vestibular system decompression sickness after an excursion to 300 m. It is concluded that the decompression table is effective in use shallower than 150 m but that the risk increases with greater depth. There is, however, only limited experience in the deeper range. There is no evidence that chamber compression with air to 10 m adversely affects decompression from deeper than 50 m. An account of the medical and physiological conditions affecting divers in these dives is given.

Decompression Sickness↗

Arteriovenous bubbles following cold water sport dives: relation to right-to-left shunting.

Neurologic injury subsequent to decompression from diving may be due to paradoxical arterialization of venous gas emboli. Of 40 divers who performed 53 open water dives after being tested for a patent foramen ovale (PFO), arterial gas emboli were detected in 7 of 13 dives, which resulted in venous bubbles. In five of these seven dives, there was evidence of a PFO by contrast transcranial Doppler sonography, indicating an increased risk of arterializing venous bubbles in divers with a PFO.

Adult↗

Factors affecting stroking patterns and body angle in diving Weddell seals under natural conditions.

Aquatic animals use a variety of strategies to reduce the energetic cost of locomotion. Efficient locomotion is particularly important for breath-holding divers because high levels of exercise may quickly deplete oxygen reserves, leading to the termination of a dive. We investigated the swimming behavior of eight adult Weddell seals, which are proficient divers, in McMurdo Sound, Antarctica. A newly developed data logger was attached to free-ranging females at their own breeding sites to record swimming speed, depth, two-dimensional accelerations (stroke frequency and body angle) and temperature. All seals conducted multiple deep dives (the mean dive depth range for each animal was 223.3+/-66.5-297.9+/-164.7 m). Prolonged gliding while descending was observed with thinner females (N=5 seals). But the fatter females (N=3 seals) exhibited only swim-and-glide swimming, characterized by intermittent stroking and fluctuating swim speed, throughout their descent and ascent. The body angles of four of the seals were restricted to less than 30 degrees by the location of breathing holes in the ice and the slope of local bathymetric features. Of these four, the three fatter seals adopted the stroke-and-glide method while the other thinner seal descended with prolonged periods of gliding. Prolonged gliding seems to be a more efficient method for locomotion because the surface time between dives of gliding seals was significantly less than that of stroking animals, despite their same stroke frequencies.

Acceleration↗

Adaptations to diving hypoxia in the heart, kidneys and splanchnic organs of harbor seals (Phoca vitulina).

Pinnipeds (seals and sea lions) have an elevated mitochondrial volume density [VV(mt)] and elevated citrate synthase (CS) and beta-hydroxyacyl-CoA dehydrogenase (HOAD) activities in their swimming muscles to maintain an aerobic, fat-based metabolism during diving. The goal of this study was to determine whether the heart, kidneys and splanchnic organs have an elevated VV(mt) and CS and HOAD activities as parallel adaptations for sustaining aerobic metabolism and normal function during hypoxia in harbor seals (Phoca vitulina). Samples of heart, liver, kidney, stomach and small intestine were taken from 10 freshly killed harbor seals and fixed in glutaraldehyde for transmission electron microscopy or frozen in liquid nitrogen for enzymatic analysis. Samples from dogs and rats were used for comparison. Within the harbor seal, the liver and stomach had the highest VV(mt). The liver also had the highest CS activity. The kidneys and heart had the highest HOAD activities, and the liver and heart had the highest lactate dehydrogenase (LDH) activities. Mitochondrial volume densities scaled to tissue-specific resting metabolic rate [VV(mt)/RMR] in the heart, liver, kidneys, stomach and small intestine of harbor seals were elevated (range 1.2-6.6x) when compared with those in the dog and/or rat. In addition, HOAD activity scaled to tissue-specific RMR in the heart and liver of harbor seals was elevated compared with that in the dog and rat (3.2x and 6.2x in the heart and 8.5x and 5.5x in the liver, respectively). These data suggest that organs such as the liver, kidneys and stomach possess a heightened ability for aerobic, fat-based metabolism during hypoxia associated with routine diving. However, a heightened LDH activity in the heart and liver indicates an adaptation for the anaerobic production of ATP on dives that exceed the animal's aerobic dive limit. Hence, the heart, liver, kidneys and gastrointestinal organs of harbor seals exhibit adaptations that promote an aerobic, fat-based metabolism under hypoxic conditions but can provide ATP anaerobically if required.

3-Hydroxyacyl CoA Dehydrogenases↗

Year-round recordings of behavioural and physiological parameters reveal the survival strategy of a poorly insulated diving endotherm during the Arctic winter.

Warm-blooded diving animals wintering in polar regions are expected to show a high degree of morphological adaptation allowing efficient thermal insulation. In stark contrast to other marine mammals and seabirds living at high latitudes, Arctic great cormorants Phalacrocorax carbo have very limited thermal insulation because of their partly permeable plumage. They nonetheless winter in Greenland, where they are exposed to very low air and water temperatures. To understand how poorly insulated diving endotherms survive the Arctic winter, we performed year-round recordings of heart rate, dive depth and abdominal temperature in male great cormorants using miniature data loggers. We also examined the body composition of individuals in the spring. Abdominal temperatures and heart rates of birds resting on land and diving showed substantial variability. However, neither hypothermia nor significantly lower heart rate levels were recorded during the winter months. Thus our data show no indication of general metabolic depression in great cormorants wintering in Greenland. Furthermore, great cormorants did not reduce their daily swimming time during the coldest months of the year to save energy; they continued to forage in sub-zero waters for over an hour every day. As birds spent extended periods in cold water and showed no signs of metabolic depression during the Arctic winter, their theoretical energy requirements were substantial. Using our field data and a published algorithm we estimated the daily food requirement of great cormorants wintering in Greenland to be 1170+/-110 g day(-1). This is twice the estimated food requirement of great cormorants wintering in Europe. Great cormorants survive the Arctic winter but we also show that they come close to starvation during the spring, with body reserves sufficient to fast for less than 3 days. Lack of body fuels was associated with drastically reduced body temperatures and heart rates in April and May. Concurrent, intense feeding activity probably allowed birds to restore body reserves. Our study is the first to record ecophysiological parameters in a polar animal on a year-round basis. It challenges the paradigm that efficient thermal insulation is a prerequisite to the colonization of polar habitats by endotherms.

Acclimatization↗

Muscle temperature and swim velocity profiles during diving in a Weddell seal, Leptonychotes weddellii.

Locomotory muscle temperature and swim velocity profiles of an adult Weddell seal were recorded over a 21 h period. The highest temperatures occurred during a prolonged surface period (mean 37.3 degrees C, S.D. 0.16 degrees C). Muscle temperature averaged 36.8 and 36.6 degrees C (S.D. 0.25 degrees C, 0.19 degrees C) during two dive bouts and showed no consistent fluctuations between dive and interdive surface intervals. Swim velocities were also constant, near 1.3 m s-1. These data indicate that past records of low aortic temperatures (35 degrees C) during and after prolonged dives are not indicative of whole-body temperature changes, and that muscle temperature, even during dives as long as 45 min, remains near 37 degrees C.

Animals↗