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Adaptations to deep breath-hold diving: respiratory and circulatory mechanics.

Respiration and circulation in diving mammals are characterized by interrelated adaptations of structure, function, and behavior that are incompletely described and understood. This speculative survey touches some of them. a) Arterial blood flow can be controlled by vasoconstriction not only in arterioles but also in large arteries. The latter physiology is not well known. b) Mechanisms that might regulate and limit nitrogen uptake are not clear, although Scholander's suggestion that airspaces become gas-free during deep dives is still accepted. c) Systemic arterial retes may be able to store oxygenated blood in some diving mammals. If so, O2 in the lung might be "skimmed off" early in a dive, leaving the N2 behind. d) Variable clusters of interdependent adaptations in diving mammals include compliant chest walls that avoid thoracic squeeze; inspiratory breath holds that maintain high lung volumes; large tidal volumes that nearly empty the lung at end-expiration (so there is near-complete turnover of lung gas with each breath); airways that are "armored" by cartilage rings all the way out to the airspaces (so that they do not close and trap gas in the lung and do permit high expiratory flow rates even at very low lung volumes); submucosal vascular retes that may prevent airway squeeze; a puzzling difference in the cross-sectional areas of trachea and bony nares; and very large lungs in shallow divers (sea otters). Study of mammalian adaptations to deep diving promises to illuminate basic issues in physiology.

Adaptation, Physiological↗

[Respiratory changes in deep diving].

Deep diving refers to saturation diving to a depth of more than 180 m (1.9 MPa ambient pressure). In the 1990s diving to 400 m may be necessary on the Norwegian continental shelf. The safety margins are narrow and at such depths the respiratory system is subject to great strain. Respiratory resistance increases and the dynamic lung volumes are reduced as the pressure increases due to enhanced gas density. Helium is used together with oxygen as breathing gas and the lower density partly normalises the dynamic lung volumes. The respiratory system imposes clear limitations on the intensity and duration of physical work during deep diving. We lack systematic studies of lung mechanics, gas exchange and respiratory regulation in the different phases of deep dives. Demonstration of possible chronic occupational respiratory diseases connected to diving is dependent on follow-up over a long time.

Adult↗

Intellectual deterioration with excessive diving (punch drunk divers).

A survey was performed on a specific occupational group of compressed air divers--the professional abalone divers of New South Wales. One aspect of this survey included the use of psychometric screening tests to elicit evidence of impaired intellectual capacity, which may be related to the compressed air diving. Results of the survey indicate that there is suggestion of intellectual impairment in almost half of this diving population. The fact that this diving group exposed themselves to much greater decompression stress than the more conventional professional diving groups suggests that these results should not be extrapolated to other diving populations. The results are supportive of the anecdotal beliefs that exist regarding this highly selective diving group, i.e., that a syndrome of reduced intellectual capacity (dementia or "punch drunkenness") may be present.

Adult↗

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↗

Sleep, mood, and fatigue during a 14-day He-O2 open-sea saturation dive to 850 fsw with excursions to 950 fsw.

To obtain information on sleep, mood, and performance of divers and surface support personnel during deep dives in the open sea, 12 divers and 12 surface support personnel were monitored during a 14-day open-sea saturation dive using the U.S. Navy Deep Diving System, Mark 2, Mod O. Divers lived in the deck decompression chambers at 850 fsw equivalent and made 5 days of excursion wet dives to approximately 950 fsw via the Personnel Transfer Capsule. Electroencephalographic and self-report measures of sleep, and measures of mood, anxiety, and 4-choice reaction time performance were obtained during a predive base-line period and throughout the dive and decompression. Results suggested that, unless personnel are rotated, there are limitations to the practical duration of very deep open-sea saturation dives caused by the accumulation of sleep debt, fatigue, and loss of psychological vigor.

Affect↗

Alveolar and peripheral venous gas changes during breath-hold diving.

Two fit young men were tested in a laboratory pool at 22 degrees C and 1 m depth before and after hyperventilation/breath-hold diving lasting 90 seconds. The percentages of alveolar oxygen and carbon dioxide were determined on a Polygas analyser and converted to partial pressures (PAO2 and PACO2) at an atmospheric pressure of 86 kPa at 20 degrees C, room temperature. PAO2 and PACO2 values were cross-checked by collecting expiratory gas in a Douglas bag and using the Haldane gas analyser. Peripheral venous blood was 'arterialized' and analysed for CO2 (PVCO2) using the Radiometer ABL1 gas analyser. For ethical reasons, arterial punctures were not considered in these healthy subjects. Peripheral venous blood was obtained under near steady-state conditions from the dorsum of the hand, which was warmed to 38 degrees C. The results of hyperventilation/breath-hold diving are illustrated. PAO2 increased after hyperventilation and decreased to below 5 kPa after the dive. PACO2 decreased after hyperventilation and increased after the dive. PVCO2 decreased after hyperventilation and increased after the dive. All values approached a common point between 3 and 5 kPa after the 90-second dive. The CO2 content of arterialized peripheral venous blood closely follows that of alveolar air.

Adolescent↗

Venous gas bubble production following cold stress during a no-decompression dive.

The effect of cold stress on venous gas bubble production was studied using Doppler ultrasonic monitoring. Ten subjects participated in four exposure regimes carried out at 78 fsw on an underwater platform for 38 min of light exercise in 10 degrees C water. Two cold exposures (1/8-in. wet suit) and two warm exposures (insulated dry suit) were each followed by rewarming in a heated bath or by endogenous heat production while insulated in a sleeping bag. Results showed that for the cold dives compared to warm dives, air consumption increased 29%, rectal temperature dropped 0.8 degrees C by the end of the dive, mean skin temperature dropped 11 degrees C, and cooling rate correlated with mean skin fold and endomorphy (P less than 0.001). A threefold increase in bubble scores (P less than 0.025) was observed following the warm dives compared to the cold dives. The results suggest that inert gas uptake is reduced as a result of peripheral vasoconstriction when the cold stress is induced at the onset of the dive and maintained throughout.

Adult↗

[Effects of variations in the ascending speed on the production of circulating gas bubbles after compressed-air diving].

Ninety-seven compressed air divers at depths of 20 to 52 msw were done. Every dive reached a tissue nitrogen saturation level greater than or equal to M value according to U.S. Navy decompression schedules and respected all prescribed decompression stop. Dives were divided in two groups according to the speed of ascent:--1st group: 33 dives (18 simulated, 15 open water) with ascent at 18 msw/min. for the first half of the distance and 10 msw/min. for the second half. No work on the bottom. Average ascent rate 14 msw/min. This profile was due to the flow limits of the outlet of our chamber during the second part of the ascent, and it was repeated in open water diving.--2nd group: 64 dives (4 simulated, 60 open water) with linear ascent at 10 msw/min. Half the open water dives were repetitive within 4 hours from the first one. Medium to heavy work on the bottom. Ultrasound Doppler bubble detection at rest and after exercise was performed at five minutes intervals and during 40 minutes after surfacing.

Diving↗

Behavior of heart rate and incidence of arrhythmia in swimming and diving.

Heart rate behaviour and the incidence of arrhythmia were recorded in 29 young subjects aged between 16 and 20 years using radiotelemetry equipment. The study consisted of four regimens: 100m freestyle swimming with and without an aqualung, up to 50m under water swimming without a breathing apparatus (skin diving) and 100m under water swimming with an aqualung (scuba diving). In the course of the swimming experiments the heart rate tracing exhibits three phases, namely: (a) a sharp rise in the first 10 s; (b) a reduction in the rate of increase, and (c) a constant plateau at approximately 184 bpm without an aqualung and 168 bpm with an aqualung. During apnoeic diving there is a slight increase in heart rate, followed by a rapid drop (vasovagal diver's bradycardia) and finally a plateau at approximately 55 bpm. Diver's bradycardia does not occur when the diver uses an aqualung; the heart rates then correspond to those observed during swimming (maximal rate approximately 169 bpm). Cardiac arrhythmia occurred in 18 instances (3 each during swimming with and without a breathing apparatus, 5 during diving with an aqualung and 7 during diving without an aqualung). 15 were cases of supraventricular extrasystoles, 12 occurred in the plateau phase. A man with a thorough endurance training exhibited in addition to supraventricular extrasystoles a transient bigeminy, substitutive AV systoles and an electric alternans during diving without an aqualung. He experienced no subjective feeling of impairment of performance, nor was there any objective reduction in efficiency.

Adolescent↗

[Acute and chronic effects of deep diving on the nervous system].

Diving deeper than 180 metres of seawater (msw) will impose neurological symptoms in most divers. Atactic signs and abnormal EEGs were found in five of 18 divers immediately after deep diving. Neuropsychological testing before and after deep diving in 64 divers revealed a reduction in autonomic reactivity (48%), increased hand tremor (27%) and impairment of spatial memory and reduced finger coordination (8%) after the dives. These results had not improved one year later. A follow-up study of 40 divers one to seven years after their last deep dive revealed that the divers experienced more problems of concentration and paresthesia in feet and hands than the controls. Two had had seizures, one had suffered episodes of transitory cerebral ischemia and one had experienced transitory global amnesia after the deep dives. In the future, oil installations at depths below 180 msw should be installed and maintained with remote control and robot technology.

Acute Disease↗

Diving decompression fails to activate complement.

The present study evaluated complement activation during decompression after air dives in a hyperbaric chamber. Intravascular bubbles were quantified by Doppler ultrasound scoring. Eighteen subjects completed 92 dives, of which 74 produced bubbles. Complement activation was assessed by plasma C3a des Arg and red-cell-bound C3d before and after each dive. These parameters of in vivo complement activation failed to show significant activation. In vitro complement activation susceptibility tests on pre-dive sera were performed to explore their association with in vivo complement activation and intravascular bubbles. Such tests failed to identify a distinct complement-sensitive group and did not correlate with in vivo complement activation during the dives and/or intravascular bubble appearance. Two subjects developed decompression sickness but were not different from the rest of the group regarding in vitro complement sensitivity or complement activation during dives.

Adult↗

Heart rate and plasma lactate responses during submerged swimming and trained diving in California sea lions, Zalophus californianus.

California sea lions, Zalophus californianus, were trained to elicit maximum voluntary breath holds during stationary underwater targeting, submerged swimming, and trained diving. Lowest heart rate during rest periods was 57 bpm. The heart rate profiles in all three protocols were dominated by a bradycardia of 20-50 bpm, and demonstrated that otariid diving heart rates were at or below resting heart rate. Venous blood samples were collected after submerged swimming periods of 1-3 min. Plasma lactate began to increase only after 2.3-min submersions. This rise in lactate and our inability to train sea lions to dive or swim submerged for periods longer than 3 min lead us to conclude that an aerobic limit had been reached. Due to the similarity of heart rate responses and swimming velocities recorded during submerged swimming and trained diving, this 2.3-min limit should approximate the aerobic dive limit in these 40-kg sea lions. Total body O2 stores, based on measurements of blood and muscle O2 stores in these animals, and prior lung O2 store analyses, were 37-43 ml O2 kg-1. The aerobic dive limit, calculated with these O2 stores and prior measurements of at-sea metabolic rates of sea lions, is 1.8-2 min, similar to that measured by the change in post-submersion lactate concentration.

Animals↗

Effects of acute and chronic baroreceptor denervation on diving responses in ducks.

To investigate the effects of barodenervation on the cardiovascular responses to forced submersion, we implanted snares around the aortic nerves innervating the baroreceptors in anesthetized (pentobarbital sodium, 35-45 mg/kg) ducks (Anas platyrhynchos). Snares were withdrawn 1 wk after surgery to effect barodenervation in the absence of complications from surgical trauma or anesthesia. After barodenervation, resting heart rate (HR), hind limb vascular resistance (HLVR), and mean arterial pressure (MAP) all increased, with different time courses, over the 16-day observation period. Bradycardia developed during forced submersion in ducks before and after barodenervation, although this response was reduced for 1-3 days after nerve section. The HR response to diving intensified 4-5 days after denervation, and the level of dive bradycardia at 16 days was similar to that before denervation. End-dive MAP fell further, as a proportion of predive MAP, with time after baroreceptor loss. The absolute level of HLVR reached during diving was unaffected by barodenervation. These results show that diving responses develop in the absence of baroreceptors, but MAP in the dive is not maintained as well in barodenervates as in baroreceptor-intact animals.

Animals↗

Evidence for nest-odour recognition in two species of diving petrel.

In nearly every procellariiform species, the sense of smell appears to be highly adapted for foraging at sea, but the sense of smell among the diving petrels is enigmatic. These birds forage at considerable depth and are not attracted to odour cues at sea. However, several procellariiform species have recently been shown to relocate their nesting burrows by scent, suggesting that these birds use an olfactory signature to identify the home burrow. We wanted to know whether diving petrels use smell in this way. We tested the common diving petrel Pelecanoides urinatrix and the South-Georgian diving petrel Pelecanoides georgicus to determine whether diving petrels were able to recognise their burrow by scent alone. To verify the efficacy of the method, we also tested a bird that is known to use olfaction for foraging and nest recognition, the thin-billed prion Pachyptila belcheri. In two-choice T-maze trials, we found that, for all species, individuals significantly preferred the odour of their own nest material to that of a conspecific. Our findings strongly suggest that an individual-specific odour provides an olfactory signature that allows burrowing petrels to recognize their own burrow. Since this ability seems to be well developed in diving petrels, our data further implicate a novel adaptation for olfaction in these two species that have been presumed to lack a well-developed sense of smell.

Animals↗

Health outcome following multi-day occupational air diving.

Acclimatization to decompression stress has been reported in caisson workers and helium-oxygen divers; however the alternative notion that the risk of decompression sickness increases with successive days of diving is widespread. We examined 201 multi-day series of 2 to 29 diving days identified retrospectively in a database of occupational air dives for evidence of acclimatization or sensitization. Decompression related health status was measured using a self-administered diver health survey; resulting scores were analyzed by linear modelling. Daily diving consisted of 1-3 dives each to mean maximum depth of 17.2 (SD 3.9) meters seawater for a mean duration of 23 (SD 17) min. Daily diver health scores increased with calculated daily risk of decompression sickness but were not influenced by the order of dives in multi-day series. Poor health outcome indicated by treated decompression sickness and diver health scores > 8 occurred early in multi-day series. There was no evidence of sensitization to decompression stress whereas the timing of poor health outcomes suggests an element of acclimatization.

Analysis of Variance↗

Venous gas emboli and complement activation after deep repetitive air diving.

Complement activity has been linked to decompression sickness (DCS), but the effects of intravascular bubbles on complement activation are poorly understood. We have investigated intravascular complement activation by measuring red blood cell (RBC)-bound C3d after repetitive air diving in man. Subjects were exposed to a single, 20 min, 170 fsw (feet of sea water) dive, or to 2 such dives with a 6-h surface interval. Doppler monitoring for venous gas emboli was performed postdive. Predive blood samples were studied to determine sensitivity of complement to activation by air bubbles. Other predive and postdive venous samples were evaluated for intravascular complement activation. No cases of DCS occurred in 39 dives. Baseline complement sensitivity appeared normally distributed, thus "sensitive" and "insensitive" subjects were not clearly distinguishable. RBC-bound C3d did not increase after 1 dive but did increase after the repetitive dive (P less than 0.05). Furthermore, maximum bubble grade was independent of complement activation.

Adult↗

Hearing threshold in sport divers: is diving really a hazard for inner ear function?

OBJECTIVE: To investigate the effect of scuba diving on the hearing threshold of sport divers who have no history of excessive noise exposure or of diving-related inner ear damage. DESIGN: Cross-sectional controlled comparison study. SETTING: General sports diving community. PARTICIPANTS: Sixty sport divers with an average of 650 dives each and at least 4 years of diving experience (mean, 10 years) were compared with a control group of 63 nondivers from our hospital staff or patients referred for rhinologic problems or benign tumors of the salivary gland. MAIN OUTCOME MEASURE: After microscopic otoscopy and tympanometry, we used pure-tone audiometry to measure the hearing threshold for air and bone conduction. The participants were divided into 3 age groups, and the hearing test results for both ears combined were statistically compared. RESULTS: There were no statistically significant differences in the hearing thresholds between sport divers and nondivers. CONCLUSIONS: The reduced hearing levels of professional divers found in other studies are probably due to the high noise levels that they have to deal with or may be a result of inner ear accidents.

Adult↗

Inner ear barotrauma in scuba divers. A long-term follow-up after continued diving.

Divers who suffer inner ear barotrauma are usually counseled to permanently avoid diving, reasoning that the injured inner ear is at increased risk of further damage. Twenty patients who suffered inner ear barotrauma while diving, but continued to dive against medical advice, were assessed on an interim basis for 1 to 12 years. As difficulty equalizing the ears during the barotraumatic event was a universal finding, prior to resuming diving, all patients were reinstructed on methods of maximizing eustachian tube function. No further deterioration of cochleovestibular function was noted. Based on these preliminary results, we conclude that recommending no further diving after inner ear barotrauma may be unnecessarily restrictive.

Adult↗