U.K. deep diving trials

Using a breathing medium of 40 kPa oxygen, remainder helium, 18 volunteer subjects participated in a series of 15 exposures to pressures equivalent to depths of 180-540 m s.w. The time of exposure at these pressures was mostly 2 days, except for the 540 m s.w. exposure, when 6 days were spent at full pressure. Compression procedures, based upon placing 'stages' at 60 m s.w. intervals, evolved with experience and proved to be a highly successful way of achieving acceptable pressure-time courses. Decompression combined slow linear release of pressure with overnight halts for sleep. On one occasion a depth of 660 m s.w. was reached by breathing 40 kPa oxygen, 10% nitrogen, remainder helium. Throughout all exposures, teams of investigators followed the changes in cardiovascular, respiratory, haematological, neurophysiological and metabolic status, and mental performance of the volunteers. Some major findings were that the neurophysiological and behavioural changes could be assigned to the motor, or vestibular, or cerebral, or autonomic systems, and were mainly first observed during compression. The subjects suffered, apparently from severe nitrogen narcosis, when breathing 10% (by volume) nitrogen in oxygen-helium at 420 m s.w. Lung ventilation was remarkably adaptable to the oxygen requirements of exercise at all depths, but cardiac output was adversely affected at 540 m s.w., particularly for heavier workloads. Ventilatory responses to carbon dioxide were significantly elevated after diving. Thermal balance was seen to be precarious, but nevertheless it was achieved by the normal subjective assessments of comfort. Water loss was affected by diminished evaporation from the skin. Skin temperature sensitivity was changed and took many days after the dives to return to normal. Energy requirements increased for work purposes, but basal metabolic rate was undisturbed. Body chemistry altered at pressures in excess of 300 m s.w., for example thyroid hormone and nitrogen balances were affected. No decompression sickness was encountered until the pressures were low, but marked haematological changes could occur during decompression. Every change that occurred during these dives reverted to normal, mostly before the end of the decompression. It is concluded that diving with oxygen-helium breathing mixtures to depths as great as 540 m s.w. can be effective and safe. An attempt is made to assess the physiological significance of the principal findings.     

Probing the limits of human deep diving

Divers breathing compressed air are restricted to 45 m depth because of the narcotic effects of nitrogen and toxic action of oxygen at increased pressures. Substitution of oxygen-helium for compressed air has permitted divers to reach 600 m. However, at depths greater than 160 m, signs and symptoms of the high pressure nervous syndrome (h.p.n.s.) occur, with tremors, myoclonic jerking, nausea, vomiting, fatigue, somnolence, e.e.g. changes, dyspnoea, and poor sleep with nightmares. It has been the objective of this Laboratory to ameliorate the symptoms of pressure-induced h.p.n.s. by the addition of small amounts of 'narcotic' nitrogen to the oxygen-helium mixture to form the Trimix breathing gas. In 1973, comparative experiments with oxygen-helium and the same divers, during compressions in only 33 min to 219.5 m and 305 m, showed such Trimix to be effective with 10% (by volume) nitrogen. Simulated dives, termed ATLANTIS, have been made with Trimix over the last 4 years to depths in excess of 610 m for 11 days, 650 m for 4 days and 686 m for 1 day. The objectives were to determine the effects of either slow or rapid rates of compression, and either 5% or 10% (by volume) nitrogen in Heliox, on the presence of h.p.n.s. or nitrogen narcosis. Measurements were made of intellectual and psychomotor performance, electrophysiological function of the brain and reflexes, lung and cardiovascular function, including arterial gas analysis at rest and work, blood chemistry and psychiatric and psychological status. The results permit the conclusion that divers may be compressed safely to depths as great as 686 m. The technique requires a slow exponential compression over days, with frequent stages lasting 14 h or more, the use of 5-8% (by volume) nitrogen in Heliox and careful selection of the divers.     

Human erythrocyte superoxide dismutase activity during deep diving

As the practical use of high pressure oxygen (HPO) in clinical medicine and the offshore industries accelerates, knowledge of its toxic nature becomes essential. In this study, divers' erythrocyte superoxide dismutase (SOD) activity was monitored during high pressure exposure and shown to decrease on average by 20% at depths greater than 150 m. Assay of total red cell SOD protein and activity established that the recorded SOD activity decrement was by loss of immuno-measurable enzyme. No evidence of intra-cellular Heinz bodies was observed. An increase of intra-membrane lipid peroxidation products, within physiological limits, was found, particularly in the denser cell fractions. Using previously in vivo pressure stressed cells, experiments at increasing O2 pressures educed that human red blood cells were oxygen "resistant" up to ten times the normal atmospheric pressure, 0.021 MPa (0.21 bar). Thereafter, a loss in SOD enzyme activity occurred with hemolysis during the in vitro decompression procedure.     

Transatlantic migration and deep mid-ocean diving by basking shark

Despite being the second largest fish, basking sharks (Cetorhinus maximus) have been assumed to remain in discrete populations. Their known distribution encompasses temperate continental shelf areas, yet until now there has been no evidence for migration across oceans or between hemispheres. Here we present results on the tracks and behaviour of two basking sharks tagged off the British Isles, one of which released its tag off Newfoundland, Canada. During the shark's transit of the North Atlantic, she travelled a horizontal distance of 9589 km and reached a record depth of 1264 m. This result provides the first evidence for a link between European and American populations and indicates that basking sharks make use of deep-water habitats beyond the shelf edge.     

Biophysical limitations on deep diving: Some limiting performance expectations

Decompression is treated according to the conventional Haldane model, but with continuously varying gas mixture, and continuous ascent. Analytical expressions are derived for the inert gas, tissue super-saturation, during dives with optimum gas mixtures. Analog computer results are used to show the supersaturation graphically, on dives of 300 ft. with 20 minutes on the bottom, and 1,000 ft. with 4 minutes on the bottom. The decompression times are much shorter than the times expected from U. S. Navy diving tables.     

Microvascular patterns in the blubber of shallow and deep diving odontocetes

Blubber, a specialized form of subdermal adipose tissue, surrounds marine mammal bodies. Typically, adipose tissue is perfused by capillaries but information on blubber vascularization is lacking. This study's goals were to: 1) describe and compare the microvasculature (capillaries, microarterioles, and microvenules) of blubber across odontocete species; 2) compare microvasculature of blubber to adipose tissue; and 3) examine relationships between blubber's lipid composition and its microvasculature. Percent microvascularity, distribution, branching pattern, and diameter of microvessels were determined from images of histochemically stained blubber sections from shallow‐diving bottlenose dolphins (Tursiops truncatus), deeper‐diving pygmy sperm whales (Kogia breviceps), deep‐diving beaked whales (Mesoplodon densirostris; Ziphius cavirostris), and the subdermal adipose tissue of domestic pigs (Sus scrofa). Tursiops blubber showed significant stratification in percent microvascularity among the superficial, middle, and deep layers and had a significantly higher percent microvascularity than all other animals analyzed, in which the microvasculature was more uniformly distributed. The percent microvasculature of Kogia blubber was lower than that of Tursiops but higher than that of beaked whales and the subdermal adipose tissue of domestic pigs. Tursiops had the most microvascular branching. Microvessel diameter was relatively uniform in all species. There were no clear patterns associating microvascular and lipid characteristics. The microvascular characteristics of the superficial layer of blubber resembled the adipose tissue of terrestrial mammals, suggesting some conservation of microvascular patterns in mammalian adipose tissue. The middle and deep layers of blubber, particularly in Tursiops, showed the greatest departure from typical mammalian microvascular arrangement. Factors such as metabolics or thermoregulation may be influencing the microvasculature in these layers. © J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Oligochaetes and water pollution in two deep Norwegian lakes

Analyses of the oligochaete fauna of two of the deepest lakes in Scandinavia — the Norwegian lakes Mjösa (450 m) and Tyrifjorden (295 m), revealed a totally different species composition in the deep profundal compared with the upper profundal - in contact with the nutrient-enriched epilimnion. In both lakes a pronounced thermal stratification develops in the summer, thus the epilimnion receiving gross organic pollution behaves differently from the profundal. The lakes are each effectively divided into two bodies of water with limited water exchange between them, i.e. one major oligotrophic body and one minor more nutrient-rich. Since the 1950s both lakes have been exposed to heavy pollution of various kinds. In Lake Mjösa in 1975 and 1976 unpleasant algal blooms of the blue-green alga Oscillatoria bornetii fa. tenuis occurred. Bottom samples obtained at the same time revealed that the deep central bottoms of the lake were totally dominated by oligotrophic oligochaete indicators, i.e. by Stylodrilus heringianus and Spirosperma ferox, while the fauna of the upper profundal in the vicinity of domestic and agricultural sewage outfalls, wood processing industries, etc. was dominated by Limnodrilus hoffmeisteri and Tubifex tubifex in great abundance, indicating enriched conditions. Several other species indicative of eutrophy, were absent, most of them belonging to the genus Potamothrix. A fairly similar situation exists in Lake Tyrifjorden, where, for instance, in the shallow bay of Steinsfjorden — heavily eutrophied by agricultural wastes — blooms of blue-green algae have caused problems from time to time. The same oligochaete communities as in Lake Mjösa distinguish the central oligotrophic bottoms from the regionally more enriched upper profundal. The likely reasons for an intact profundal oligochaete fauna are great volumes of oxygen-rich hypolimnic water of low temperature and a high bottom/lake surface area ratio.     

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.     

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.     

Foraging in the darkness of the southern ocean: influence of bioluminescence on a deep diving predator

How non-echolocating deep diving marine predators locate their prey while foraging remains mostly unknown. Female southern elephant seals (SES) (Mirounga leonina) have vision adapted to low intensity light with a peak sensitivity at 485 nm. This matches the wavelength of bioluminescence produced by a large range of marine organisms including myctophid fish, SES's main prey. In this study, we investigated whether bioluminescence provides an accurate estimate of prey occurrence for SES. To do so, four SES were satellite-tracked during their post-breeding foraging trip and were equipped with Time-Depth-Recorders that also recorded light levels every two seconds. A total of 3386 dives were processed through a light-treatment model that detected light events higher than ambient level, i.e. bioluminescence events. The number of bioluminescence events was related to an index of foraging intensity for SES dives deep enough to avoid the influence of natural ambient light. The occurrence of bioluminescence was found to be negatively related to depth both at night and day. Foraging intensity was also positively related to bioluminescence both during day and night. This result suggests that bioluminescence likely provides SES with valuable indications of prey occurrence and might be a key element in predator-prey interactions in deep-dark marine environments.     

Stroke and glide of wing-propelled divers: deep diving seabirds adjust surge frequency to buoyancy change with depth

In order to increase locomotor efficiency, breath-holding divers are expected to adjust their forward thrusts in relation to changes of buoyancy with depth. Wing propulsion during deep diving by Brünnich's guillemots (Uria lomvia) was measured in the wild by high-speed (32 Hz) sampling of surge (tail-to-head) and heave (ventral-to-dorsal) accelerations with bird-borne data loggers. At the start of descent, the birds produced frequent surges (3.2 Hz) during both the upstroke and the downstroke against buoyancy to attain a mean speed of 1.2-1.8 m s(-1) that was close to the expected optimal swim speed. As they descended deeper, the birds decreased the frequency of surges to 2.4 Hz, relaying only on the downstroke. During their ascent, they stopped stroking at 18 m depth, after which the swim speed increased to 2.3 m s(-1), possibly because of increasing buoyancy as air volumes expanded. This smooth change of surge frequency was achieved while maintaining a constant stroke duration (0.4-0.5 s), presumably allowing efficient muscle contraction.     

Locomotor muscle profile of a deep (Kogia breviceps) versus shallow (Tursiops truncatus) diving cetacean

When a marine mammal dives, breathing and locomotion are mechanically uncoupled, and its locomotor muscle must power swimming when oxygen is limited. The morphology of that muscle provides insight into both its oxygen storage capacity and its rate of oxygen consumption. This study investigated the m. longissimus dorsi, an epaxial swimming muscle, in the long duration, deep‐diving pygmy sperm whale (Kogia breviceps) and the short duration, shallow‐diving Atlantic bottlenose dolphin (Tursiops truncatus). Muscle myoglobin content, fiber type profile (based upon myosin ATPase and succinate dehydrogenase assays), and fiber size were measured for five adult specimens of each species. In addition, a photometric analysis of sections stained for succinate dehydrogenase was used to create an index of mitochondrial density. The m. longissimus dorsi of K. breviceps displayed significantly a) higher myoglobin content, b) larger proportion of Type I (slow oxidative) fibers by area, c) larger mean fiber diameters, and d) lower indices of mitochondrial density than that of T. truncatus. Thus, this primary swimming muscle of K. breviceps has greater oxygen storage capacity, reduced ATP demand, and likely a reduced rate of oxygen consumption relative to that of T. truncatus. The locomotor muscle of K. breviceps appears able to ration its high onboard oxygen stores, a feature that may allow this species to conduct relatively long duration, deep dives aerobically. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

An application of optimal diving models to diving behaviour of Brünnich's guillemots

Although theoretical models predict that the quality of foraging patches has little effect on optimal dive time with increasing depth, many empirical studies show that dive time at a given depth may vary. We developed a model that incorporated patch quality as a parameter of energy intake as a nonlinear function of time, and applied it to the diving behaviour of Brünnich's guillemots, Uria lomvia. The model indicated that optimal dive time can vary widely depending on the parameter. It also explained the convergence of observed dive times with travel time. Assuming the birds dived optimally, this parameter can be estimated from travel time and dive time for each dive. Foraging patches with larger estimated parameter values were favoured by the birds, suggesting that the parameter indicated patch quality. We used this parameter to test an optimal patch use model in divers. The results indicate that Brünnich's guillemots adjust their diving behaviour adaptively depending on patch quality, and that the optimal diving model is valid for prediction of observed dive patterns if patch quality is incorporated appropriately. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Microvascular characteristics of the acoustic fats: Novel data suggesting taxonomic differences between deep and shallow‐diving odontocetes

Odontocetes have specialized mandibular fats, the extramandibular (EMFB) and intramandibular fat bodies (IMFB), which function as acoustic organs, receiving and channeling sound to the ear during hearing and echolocation. Recent strandings of beaked whales suggest that these fat bodies are susceptible to nitrogen (N₂) gas embolism and empirical evidence has shown that the N₂ solubility of these fat bodies is higher than that of blubber. Since N₂ gas will diffuse from blood into tissue at any blood/tissue interface and potentially form gas bubbles upon decompression, it is imperative to understand the extent of microvascularity in these specialized acoustic fats so that risk of embolism formation when diving can be estimated. Microvascular density was determined in the EMFB, IMFB, and blubber from 11 species representing three odontocete families. In all cases, the acoustic tissues had less (typically 1/3 to 1/2) microvasculature than did blubber, suggesting that capillary density in the acoustic tissues may be more constrained than in the blubber. However, even within these constraints there were clear phylogenetic differences. Ziphiid (Mesoplodon and Ziphius, 0.9 ± 0.4% and 0.7 ± 0.3% for EMFB and IMFB, respectively) and Kogiid families (1.2 ± 0.2% and 1.0 ± 0.01% for EMFB and IMFB, respectively) had significantly lower mean microvascular densities in the acoustic fats compared to the Delphinid species (Tursiops, Grampus, Stenella, and Globicephala, 1.3 ± 0.3% and 1.3 ± 0.3% for EMFB and IMFB, respectively). Overall, deep‐diving beaked whales had less microvascularity in both mandibular fats and blubber compared to the shallow‐diving Delphinids, which might suggest that there are differences in the N₂ dynamics associated with diving regime, phylogeny, and tissue type. These novel data should be incorporated into diving physiology models to further understand potential functional disruption of the acoustic tissues due to changes in normal diving behavior.