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Gas transport and blood acid-base balance in diving sea snakes.

The values of hemoglobin concentration, Hb-O2 affinity and buffering capacity of the blood of six sea snake species considerably overlap values from terrestrial squamates. Decreased blood pH had little effect on the P50 but increased the n-value of Hb-O2 equilibrium curves. The O2 saturation of blood in the dorsal aorta varied between about 30 and 70% during voluntary diving in Acalyptophis peronii and Lapemis hardwickii. Voluntary dives ended when the lung PP02 was about 50 mm Hg and the arterial PO2 about 30 mm Hg indicating that roughly half of the O2 reserves had been used. In conjunction with relatively stable blood lactate concentration and pH, this indicates that voluntary dives occurred largely aerobically. In contrast, forced dives resulted in depletion of O2 reserves and large changes in blood acid-base balance. Long recovery periods following forced dives are inconsistent with field observations and thus suggest that extensive anaerobic metabolism does not normally occur in sea snakes. Bradycardia was not evident during forced dives. Large differences in PO2 between the lung and dorsal aorta indicated considerable right to left shunting either in the heart or in the lung. Venous blood represented over 50% of the systemic flow when there was considerable O2 in the lung. Therefore blood PO2 may remain relatively low despite elevated lung PO2 resulting from diving. In view of substantial capability for extra-pulmonary gas exchange, high shunting reduces the possibility of losing O2 through the skin and also may help prevent decompression sickness following deep dives.

Acid-Base Equilibrium↗

Blood-respiratory and acid-base changes during extended diving in the bimodally respiring freshwater turtle Rheodytes leukops.

Changes in blood-gas, acid-base, and plasma-ion status were investigated in the bimodally respiring turtle, Rheodytes leukops, during prolonged dives of up to 12 h. Given that R. leukops routinely submerges for several hours, the objective of this study was to determine whether voluntarily diving turtles remain aerobic and simultaneously avoid hypercapnic conditions over increasing dive lengths. Blood PO(2), PCO(2), and pH, as well as plasma concentrations of lactate, glucose, Na(+), K(+), Cl(-), total Ca, and total Mg were determined in venous blood collected from the occipital sinus. Blood PO(2) declined significantly with dive length; however, oxy-haemoglobin saturation remained greater than 30% for all R. leukops sampled. No changes were observed in blood PCO(2), pH, [HCO(3)(-)], or plasma glucose, with increasing dive length. Despite repeated dives lasting more than 2 h, plasma lactate remained less than 3 mmol l(-1) for all R. leukops sampled, indicating the absence of anaerobiosis. Compensatory acid-base adjustments associated with anaerobiosis (e.g. declining [Cl(-)], increasing total [Ca] and [Mg]) were likewise absent, with plasma-ion concentrations remaining stable with increasing dive length. Results indicate that R. leukops utilises aquatic respiration to remain aerobic during prolonged dives, thus effectively avoiding the development of a metabolic and respiratory acidosis.

Acid-Base Equilibrium↗

Mechanical impedance of the respiratory tract in divers before and after simulated deep dives.

Previous studies have inconsistently shown changes in expiratory flows and volumes as well as diffusion capacity of the lungs after single dives and several diving related occupational conditions were considered as possible underlying factors. In this study mechanical impedance of the airways was measured before and after simulated dives to non-invasively determine whether there is evidence for lung function impairment due to hyperbaric exposure. Thirty-three healthy male divers employing air self-contained underwater breathing apparatus were randomly assigned to dry and wet chamber dives in a cross-over design to 600 kPa ambient pressure (total duration 43 min, bottom time 15 min, water temperature 24 degrees C). Immediately before and after diving, oscillometric parameters-e. g. resistance and reactance of the respiratory tract-were measured at defined frequencies (5, 20 Hz). Spirometry was carried out as well (FVC, FEV(1), MEF 25-75). No significant changes between post-exposure values and baseline values were detected by respiratory impedance and spirometry. Diving in accordance to diving regulations and without excessive workload is not a source for acute obstructive lung function changes as the obtained oscillometric data suggested. Moreover this study could not confirm changes in spirometry after simulated diving exposure.

Adult↗

No changes in lung function after a saturation dive to 2.5 MPa with intermittent reduction in Po2 during decompression.

Decompression stress and exposure to hyperoxia may cause a reduction in transfer factor of the lung for carbon monoxide and in maximal aerobic capacity after deep saturation dives. In this study lung function and exercise capacity were assessed before and after a helium-oxygen saturation dive to a pressure of 2.5 MPa where the decompression rate was reduced compared with previous deep dives, and the hyperoxic exposure was reduced by administering oxygen intermittently at pressures of 50 and 30 kPa during decompression. Eight experienced divers of median age 41 years (range 29-48) participated in the dive. The incidence of venous gas microemboli was low compared with previous deep dives. Except for one subject having treatment for decompression sickness, no changes in lung function or angiotensin converting enzyme, a marker of pulmonary endothelial cell damage, were demonstrated. The modified diving procedures with respect to decompression rate and hyperoxic exposure may have contributed to the lack of changes in lung function in this dive compared with previous deep saturation dives.

Adult↗

Optimal diving under the risk of predation.

Many air-breathing aquatic foragers may be killed by aerial or subsurface predators while recovering oxygen at the surface; yet the influence of predation risk on time allocation during dive cycles is little known in spite of numerous studies on optimal diving. We modeled diving behavior under the risk of predation at the surface. The relationship between time spent at the surface and the risk of death is predicted to influence the optimal surface interval, regardless of whether foragers accumulate energy at a constant rate while at the food patch, deplete food resources over the course of the dive, or must search for food during the dive. When instantaneous predation risk during a single surface interval decreases with time spent at the surface, a diver should increase its surface interval relative to that which maximizes energy intake, thereby increasing dive durations and reducing the number of surfacings per foraging bout. When instantaneous risk over a single surface interval does not change or increases with increasing time at the surface, divers should decrease their surface interval (and consequently their dive duration) relative to that which maximizes energy intake resulting in more dives per foraging bout. The fitness consequences of selecting a suboptimal surface interval vary with the risk function and the way divers harvest energy when at depth. Finally, predation risk during surface intervals should have important consequences for habitat selection and other aspects of the behavioral ecology of air-breathing aquatic organisms.

Animals↗

Acute otitis externa in divers working in the North Sea: a microbiological survey of seven saturation dives.

Saturation diving is an important and widely used technique in the Offshore Oil Industry. During 1974-5 two saturation dives in the North Sea were terminated because of outbreaks of incapacitating otitis externa, and others were disrupted. Pseudomonas aeruginosa was consistently isolated from the ears of affected divers. Because complex work schedules were threatened seven subsequent dives were subjected to microbiological monitoring and control. Colonization of ear canal with P. aeruginosa or with other gram-negative bacilli occurred in 39 (67%) of the 58 divers studied, usually within 7 days of starting the dive. Data obtained by serotyping this isolations of P. aeruginosa suggested that a single infected diver may be the source of organisms which rapidly spread to his colleagues and throughout the living chambers, that the living chambers may constitute a reservoir of infection during and between dives, and that certain serotypes of P. aeruginosa are more likely than others to colonize the ear canal in the conditions of a saturation dive. The control measures used during the dives were only partially effective, but none of the divers suffered severe pain and all the dives were an operational success.

Corynebacterium↗

Role of cardiorespiratory abnormalities, smoking and dive characteristics in the manifestations of neurological decompression illness.

1. Blind analysis of contrast echocardiograms to detect intracardiac shunts, blind analysis of lung function tests for evidence of small airways disease, smoking history and dive characteristics were examined in an attempt to explain neurological symptoms that occurred within 5 min of surfacing from unprovocative dives. 2. Pulmonary abnormalities were significantly more frequent in those divers without intracardiac shunts (50%) than in those with shunts (0%). Smoking was more common in those divers without shunts (55% versus 15%), although this just failed to reach conventional significance levels. Divers without shunts experienced cerebral rather than spinal symptoms after significantly shallower dives with lower tissue nitrogen loads. Depths of dives, tissue nitrogen loads and clinical manifestations in those divers without shunts were similar to the findings in divers who had symptoms after rapid ascents. Despite conservative dive profiles, clinical manifestations in divers with shunts resembled those observed after missed decompression stops. 3. The findings suggest that occult lung disease, and probably smoking, increase the risk of neurological symptoms, even after unprovocative dives, and the similarity of the dive profiles and clinical manifestations to cases with rapid ascents suggest that pulmonary barotrauma and arterial gas embolism are responsible. In divers with intracardiac shunts the different dive profiles and clinical manifestations imply that there is another mechanism, involving different tissue and bubble nitrogen kinetics resulting in venous gas liberation and peripheral amplification in embolized tissues, rather than paradoxical embolism per se.

Adult↗

Decompression sickness following breath-hold diving.

Despite convincing evidence of a relationship between breath-hold diving and decompression sickness (DCS), the causal connection is only slowly being accepted. Only the more recent textbooks have acknowledged the risks of repetitive breath-hold diving. We compare four groups of breath-hold divers: (1) Japanese and Korean amas and other divers from the Pacific area, (2) instructors at naval training facilities, (3) spear fishers, and (4) free-dive athletes. While the number of amas is likely decreasing, and Scandinavian Navy training facilities recorded only a few accidents, the number of spear fishers suffering accidents is on the rise, in particular during championships or using scooters. Finally, national and international associations (e.g., International Association of Free Drives [IAFD] or Association Internationale pour Le Developpment De L'Apnee [AIDA]) promote free-diving championships including deep diving categories such as constant weight, variable weight, and no limit. A number of free-diving athletes, training for or participating in competitions, are increasingly accident prone as the world record is presently set at a depth of 171 m. This review presents data found after searching Medline and ISI Web of Science and using appropriate Internet search engines (e.g., Google). We report some 90 cases in which DCS occurred after repetitive breath-hold dives. Even today, the risk of suffering from DCS after repetitive breath-hold diving is often not acknowledged. We strongly suggest that breath-hold divers and their advisors and physicians be made aware of the possibility of DCS and of the appropriate therapeutic measures to be taken when DCS is suspected. Because the risk of suffering from DCS increases depending on depth, bottom time, rate of ascent, and duration of surface intervals, some approaches to assess the risks are presented. Regrettably, none of these approaches is widely accepted. We propose therefore the development of easily manageable algorithms for the prevention of those avoidable accidents.

Decompression Sickness↗

The autonomic nervous control of heart rate in ducks during voluntary diving.

Autonomic nervous control of heart rate was studied in voluntarily diving ducks (Aythya affinis). Ducks were injected with the muscarinic blocker atropine, the beta-adrenergic blocker nadolol, the beta-adrenergic agonist isoproterenol, and a combination of both atropine and nadolol. Saline injection was used as a control treatment. The reduction in heart rate (from the predive level) normally seen during a dive was abolished by atropine. Nadolol reduced heart rate during all phases of diving activity-predive, dive, and postdive-indicating that sympathetic output to the heart was not withdrawn during diving. Isoproterenol increased heart rate before, during, and after the dive, although the proportional increase in heart rate was not as high during the dive as compared with the increase in routine heart rate or heart rate during the predive or postdive phase. The parasympathetic system predominates in the control of heart rate during diving despite the maintenance of efferent sympathetic influences to the heart, perhaps due to accentuated antagonism between the two branches of the autonomic nervous system.

Animals↗

Metabolic limits on dive duration and swimming speed in the southern elephant seal Mirounga leonina.

The ability of air-breathing marine predators to forage successfully depends on their ability to remain submerged. This is in turn related to their total O(2) stores and the rate at which these stores are used up while submerged. Body size was positively related to dive duration in a sample of 34 adult female southern elephant seals from Macquarie Island. However, there was no relationship between body size and dive depth. This indicates that smaller seals, with smaller total O(2) stores, make shorter dives than larger individuals but operate at similar depths, resulting in less time being spent at depth. Nine adult female elephant seals were also equipped with velocity time depth recorders. In eight of these seals, a plot of swimming speed against dive duration revealed a cloud of points with a clear upper boundary. This boundary could be described using regression analysis and gave a significant negative relationship in most cases. These results indicate that metabolic rate varies with activity levels, as indicated by swimming speed, and that there are quantifiable limits to the distance that a seal can travel on a dive of a given swimming speed. However, the seals rarely dive to these physiological limits, and the majority of their dives are well within their aerobic capacity. Elephant seals therefore appear to dive in a way that ensures that they have a reserve of O(2) available.

Aerobiosis↗

Diving behaviour of a reptile (Crocodylus johnstoni) in the wild: interactions with heart rate and body temperature.

The differences in physical properties of air and water pose unique behavioural and physiological demands on semiaquatic animals. The aim of this study was to describe the diving behaviour of the freshwater crocodile Crocodylus johnstoni in the wild and to assess the relationships between diving, body temperature, and heart rate. Time-depth recorders, temperature-sensitive radio transmitters, and heart rate transmitters were deployed on each of six C. johnstoni (4.0-26.5 kg), and data were obtained from five animals. Crocodiles showed the greatest diving activity in the morning (0600-1200 hours) and were least active at night, remaining at the water surface. Surprisingly, activity pattern was asynchronous with thermoregulation, and activity was correlated to light rather than to body temperature. Nonetheless, crocodiles thermoregulated and showed a typical heart rate hysteresis pattern (heart rate during heating greater than heart rate during cooling) in response to heating and cooling. Additionally, dive length decreased with increasing body temperature. Maximum diving length was 119.6 min, but the greatest proportion of diving time was spent on relatively short (<45 min) and shallow (<0.4 m) dives. A bradycardia was observed during diving, although heart rate during submergence was only 12% lower than when animals were at the surface.

Activity Cycles↗

Body cooling and its energetic implications for feeding and diving of tufted ducks.

Wintering in a temperate climate with low water temperatures is energetically expensive for diving ducks. The energy costs associated with body cooling due to diving and ingesting large amounts of cold food were measured in tufted ducks (Aythya fuligula) feeding on zebra mussels (Dreissena polymorpha), using implanted heart rate and body temperature transmitters. The effects of diving depth and food ingestion were measured in two sets of experiments: we measured body cooling and energy costs of six tufted ducks diving to different depths in a 6-m-deep indoor tank; the costs for food ingestion and crushing mussel shells were assessed under seminatural winter conditions with the same ducks feeding on mussels in a 1.5-m-deep outdoor pond. Body temperature dropped during feeding bouts and increased gradually during intermittent resting periods. The temperature drop increased linearly with dive duration. The rate of body cooling increased with feeding depth, but it was lower again at depths below 4 m. Half of the increment in energy costs of diving can be attributed to thermoregulatory heat production, of which approximately 50% is generated after diving to warm up the body. The excess costs for ducks feeding on large-sized mussels could be entirely explained by the estimated energy cost necessary to compensate the heat loss following food ingestion, suggesting that the heat production from shell crushing substituted for thermoregulation. Recovery from heat loss is probably a major component of the activity budget of wintering diving ducks.

Adaptation, Physiological↗

Diving: what to tell the patient with asthma and why?

Until a decade ago, divers with asthma were uniformly barred from diving with compressed air. This prohibition was based more on theoretical concerns for barotrauma than on actual data. Follow-up studies, although retrospective, do not support a ban on recreational or commercial diving for divers with stable asthma. These studies have noted that, despite the prohibition on diving, many divers with asthma have logged multiple dives without negative consequences. When those who have suffered diving-related barotrauma have undergone physiologic testing, measurements of small airways dysfunction (maximal mid-expiratory flow rates) have been lower than measurements for comparable divers who have never suffered diving accidents. Follow-up studies with long-term commercial divers have shown that a small percentage of individuals who have sufferred diving-related barotrauma also develop abnormal maximal mid-expiratory flow rates and even some airway hyperreactivity. These latter findings correlate with the changes that occur in chronic asthmatic patients, especially those who are not well treated. The decision as to whether an asthmatic patient should be allowed to dive rests on the individual's physiologic function, maturity, and insight into the consequences of poorly managed airway inflammation and bronchospasm.

Asthma↗

Ascent exhalations of Antarctic fur seals: a behavioural adaptation for breath-hold diving?

Novel observations collected from video, acoustic and conductivity sensors showed that Antarctic fur seals consistently exhale during the last 50-85% of ascent from all dives (10-160 m, n > 8000 dives from 50 seals). The depth of initial bubble emission was best predicted by maximum dive depth, suggesting an underlying physical mechanism. Bubble sound intensity recorded from one seal followed predictions of a simple model based on venting expanding lung air with decreasing pressure. Comparison of air release between dives, together with lack of variation in intensity of thrusting movement during initial descent regardless of ultimate dive depth, suggested that inhaled diving lung volume was constant for all dives. The thrusting intensity in the final phase of ascent was greater for dives in which ascent exhalation began at a greater depth, suggesting an energetic cost to this behaviour, probably as a result of loss of buoyancy from reduced lung volume. These results suggest that fur seals descend with full lung air stores, and thus face the physiological consequences of pressure at depth. We suggest that these regular and predictable ascent exhalations could function to reduce the potential for a precipitous drop in blood oxygen that would result in shallow-water blackout.

Adaptation, Physiological↗

Haematology studies during a 350-metre dive.

Routine haematological monitoring of 6 deep-sea divers was performed pre-dive, during the three phases of the 350 m working dive, and at two post-dive medical examinations. In the compression phase a small percentage (less than 5%) of each subject's red cells became non-discoid in shape and this trend continued during the 6 d at 350 m. Concomitantly each subject was mildly dehydrated by compression diuresis and had a raised haematocrit (+5%); all other haematological parameters remained within normal limits. The number of morphologically aberrant cells continued to increase during decompression but were not present at the 1 month post-dive medical examination. The subjects' mean relative reticulocyte number was decreased during the dive to 0.4%, showing a rapid and sustained increase to 2.1% at both post-dive examinations. The red cell count was reduced by 10% during the course of the dive. Hb concentration and haematocrit evidenced no differences between the pre- or post-dive measurements.

Adult↗

Aerobic exercise before diving reduces venous gas bubble formation in humans.

We have previously shown in a rat model that a single bout of high-intensity aerobic exercise 20 h before a simulated dive reduces bubble formation and after the dive protects from lethal decompression sickness. The present study investigated the importance of these findings in man. Twelve healthy male divers were compressed in a hyperbaric chamber to 280 kPa at a rate of 100 kPa min(-1) breathing air and remaining at pressure for 80 min. The ascent rate was 9 m min(-1) with a 7 min stop at 130 kPa. Each diver underwent two randomly assigned simulated dives, with or without preceding exercise. A single interval exercise performed 24h before the dive consisted of treadmill running at 90% of maximum heart rate for 3 min, followed by exercise at 50% of maximum heart rate for 2 min; this was repeated eight times for a total exercise period of 40 min. Venous gas bubbles were monitored with an ultrasonic scanner every 20 min for 80 min after reaching surface pressure. The study demonstrated that a single bout of strenuous exercise 24h before a dive to 18 m of seawater significantly reduced the average number of bubbles in the pulmonary artery from 0.98 to 0.22 bubbles cm(-2)(P= 0.006) compared to dives without preceding exercise. The maximum bubble grade was decreased from 3 to 1.5 (P= 0.002) by pre-dive exercise, thereby increasing safety. This is the first report to indicate that pre-dive exercise may form the basis for a new way of preventing serious decompression sickness.

Adult↗

Three year follow up of a self certification system for the assessment of fitness to dive in Scotland.

BACKGROUND: The need for routine medical examinations of sport divers in the Scottish Sub-Aqua Club (Scot-SAC) was revised in March 2000, and a new system using a self administered screening questionnaire was developed to allow divers to be assessed when necessary by doctors with diving medicine experience. OBJECTIVE: To assess the effect of the new medical system on medical referee workload, diver exclusion rates, and diving incident frequency. METHODS: All divers were required to complete a questionnaire to screen for conditions that might affect fitness to dive. Divers answering "Yes" to any of the questions had their medical background assessed by a diving doctor, and, if necessary, received a clinical examination or investigation. The rate of diver exclusions based on the questionnaire response was recorded in conjunction with analysis of the incident reports. RESULTS: The number of forms requiring review by diving doctors increased from 1.2% to 5.7% (p<0.0001, 95% confidence interval (CI) -0.06 to -0.03) in the year after the introduction of the new medical system and gradually increased in subsequent years to 7.7% (p<0.0001, 95% CI -0.08 to -0.05). The number of divers failing to be certified fit to dive increased slightly from 0.7% to 1.0% after one year (p = 0.26, 95% CI -0.01 to 0.00) and subsequently to 2.0% (p = 0.0003, 95% CI 0.02 to -0.01) after three years. Most divers were certified fit to dive on the basis of the questionnaire alone, and only 0.9% required objective investigation (such as exercise testing or echocardiography). Analysis of the incidents during three years of follow up confirmed that no incident occurred because of an undetected pre-existing medical condition. Two incidents involved divers with hypertension, but both had received medical examinations and investigation based on their responses to the questionnaire. CONCLUSION: The new self administered questionnaire system appears to be an effective screening tool for the detection of divers requiring detailed assessment by doctors with diving medicine experience.

Certification↗

Inhibition of shivering in hypothermic seals during diving.

The mammalian response to hypothermia is increased metabolic heat production, usually by way of muscular activity, such as shivering. Seals, however, have been reported to respond to diving with hypothermia, which in other mammals under other circumstances would have elicited vigorous shivering. In the diving situation, shivering could be counterproductive, because it obviously would increase oxygen consumption and therefore reduce diving capacity. We have measured the electromyographic (EMG) activity of three different muscles and the rectal and brain temperature of hooded seals (Cystophora cristata) while they were exposed to low ambient temperatures in a climatic chamber and while they performed a series of experimental dives in cold water. In air, the seals had a normal mammalian shivering response to cold. Muscles were recruited in a sequential manner until body temperature stopped dropping. Shivering was initiated when rectal temperature fell below 35.3 +/- 0.6 degrees C (n = 6). In the hypothermic diving seal, however, the EMG activity in all of the muscles that had been shivering vigorously before submergence was much reduced, or stopped altogether, whereas it increased again upon emergence but was again reduced if diving was repeated. We conclude that shivering is inhibited during diving to allow a decrease in body temperature whereby oxygen consumption is decreased and diving capacity is extended.

Air↗