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Depth-dependent acoustic features of diving sperm whales (Physeter macrocephalus) in the Gulf of Mexico.

Three-dimensional dive trajectories of three sperm whales in the Gulf of Mexico have been obtained by measuring the relative arrival times and bearings of the animals' acoustic multipath reflections, using two elements of a towed hydrophone array deployed at an unknown depth and orientation. Within the first 6-12 min of the start of a dive, the intervals between successive "clicks" of all three whales corresponded closely with the two-way travel time of an acoustic pulse traveling vertically between the animals' position and the ocean bottom. The click spectra contained multiple peaks, including a faint band of energy originally centered near 10 kHz. As the animals descended over 500 m in depth, the center frequency of this band shifted to nearly 15 kHz, but subsequently remained near this value during the rest of the dive. This frequency shift is consistent with that expected from energy scattering from an ensemble of incompressible small-scale air-filled resonators, with diameters on the order of 4 mm. One possible candidate for such an ensemble is proposed to reside in the collapsed frontal sac of the animal. A comparison of the received levels for the bottom and direct multipath arrivals indicates that the whales' acoustic directivity must range between 10-30 dB in the 5-20-kHz region.

Acoustics↗

Dissociation of bradycardia and arterial constriction during diving in the seal Phoca vitulina.

Bradycardia associated with diving in the harbor seal has been dissociated from the arterial constrictor response by intracardiac pacing. Development of arterial constriction does not depend upon the development of bradycardia. During pacing, arterial constriction continues in the absence of bradycardia. Increases in heart rate to values greater than 120 beats per minute during a dive produce a progressive decrease in mean aortic pressure, which suggests that one major function of bradycardia is to reduce cardiac output, thus matching left ventricular output to the restricted vascular bed and decreased venous return associated with diving.

Animals↗

Dolphin lung collapse and intramuscular circulation during free diving: evidence from nitrogen washout.

Intramuscular nitrogen tensions in Tursiops truncatus after a schedule of repetitive ocean dives suggest a lung collapse depth of about 70 meters and suggest that intramuscular circulation is maintained during unrestrained diving in the open ocean. Therefore, the bottle-nosed dolphin is not protected by lung collapse from the decompression hazards of dives to depths shallower than 70 meters.

Adipose Tissue↗

Diving depths and energy requirements of king penguins.

During 4- to 8-day periods at sea, half of 2595 dives of three king penguins were more than 50 meters and two dives exceeded 240 meters. The at-sea metabolic rate, estimated from the turnover of tritiated water, was 2.8 times the standard metabolic rate and requires about 2.5 kilograms of squid per day. Ten percent or less of the dives may result in prey capture.

Animals↗

Self-reported long-term effects of diving and decompression illness in recreational scuba divers.

The aim of this study was to examine the long-term effects of neurological decompression illness (NDCI) on recreational divers. Thirty-seven divers who had been treated for neurological decompression illness at least 2 years previously, and a control group of 50 divers with no history of decompression illness, responded to a postal questionnaire. Divers in the accident group reported more symptoms of neurological damage, were more likely to believe that diving had a deleterious effect on their health and were more likely to indicate symptoms of psychiatric morbidity. The quantity of diving in the year preceding the survey was associated with reports of neurological damage in both groups and with symptoms of psychiatric morbidity in the accident group. Although requiring confirmation from a longitudinal study, these results suggest that recreational diving can have negative long-term consequences for health, particularly after decompression illness.

Adult↗

Relevance of postmortem radiology to the diagnosis of fatal cerebral gas embolism from compressed air diving.

AIMS: To test the hypothesis that artefact caused by postmortem off-gassing is at least partly responsible for the presence of gas within the vascular system and tissues of the cadaver following death associated with compressed air diving. METHODS: Controlled experiment sacrificing sheep after a period of simulated diving in a hyperbaric chamber and carrying out sequential postmortem computed tomography (CT) on the cadavers. RESULTS: All the subject sheep developed significant quantities of gas in the vascular system within 24 hours, as demonstrated by CT and necropsy, while the control animals did not. CONCLUSIONS: The presence of gas in the vascular system of human cadavers following diving associated fatalities is to be expected, and is not necessarily connected with gas embolism following pulmonary barotrauma, as has previously been claimed.

Animals↗

Effect of commercial diving on ventilatory function.

A retrospective analysis of spirometric data from divers attending for annual medical examination at intervals from three to nine years was carried out to examine the long term effect of diving on lung volumes. Those divers with records over a three or four year interval (group 1, n = 224) showed a mean reduction of forced vital capacity (FVC) of 240 ml; those with records over a five or more years interval (group 2, n = 123) showed a reduction of FVC of 400 ml. These reductions remained significant when expressed as a percentage of predicted normal values. The reduction of FVC between records did not correlate with the diver's age, maximum operating depth, duration of diving career, or weight change but was positively correlated with the initial FVC. The reductions in FVC were similar in smokers and non-smokers. The change in forced expiratory volume in one second (FEV1) followed a similar pattern but was less pronounced than the effect on FVC. The decline in FVC associated with diving occurs from values of FVC that are above the predicted normal; few values below predicted normal were observed. The effect may represent either a gradual return towards the predicted normal or a pathological reduction in lung volume.

Adult↗

Influence of occupational diving upon the nervous system: an epidemiological study.

Neurological signs and symptoms were recorded from 156 air and saturation divers and 100 controls. Fifty one (33%) of the divers had had symptoms from the central nervous system during decompression. Also, 22 (14%) had been unconscious while diving. In total 79 (51%) had had decompression sickness (DCS). Twelve (8%) of the divers and no controls had had specific neurological symptoms (vision disturbances, vertigo, reduced skin sensitivity) in non-diving situations, and six (4%) of the divers (no controls) had had episodes of cerebral dysfunction (seizures, transient cerebral ischaemia, transient amnesia). The divers had significantly more general symptoms from the nervous system and more abnormal neurological findings than the controls. The most prominent symptoms were difficulties in concentration and problems with long and short term memory. The most prominent abnormal findings in the divers were compatible with dysfunction in the distal spinal cord or nerve roots, and polyneuropathy. The general neurological symptoms and findings were independently significantly correlated with diving exposure, prevalence of DCS, and age.

Adult↗

Diving and the risk of barotrauma.

STUDY OBJECTIVES: Pulmonary barotrauma (PBT) of ascent is a feared complication in compressed air diving. Although certain respiratory conditions are thought to increase the risk of suffering PBT and thus should preclude diving, in most cases of PBT, risk factors are described as not being present. The purpose of our study was to evaluate factors that possibly cause PBT. DESIGN: We analyzed 15 consecutive cases of PBT with respect to dive factors, clinical and radiologic features, and lung function. They were compared with 15 cases of decompression sickness without PBT, which appeared in the same period. RESULTS: Clinical features of PBT were arterial gas embolism (n = 13), mediastinal emphysema (n = 1), and pneumothorax (n = 1). CT of the chest (performed in 12 cases) revealed subpleural emphysematous blebs in 5 cases that were not detected in preinjury and postinjury chest radiographs. A comparison of predive lung function between groups showed significantly lower midexpiratory flow rates at 50% and 25% of vital capacity in PBT patients (p < 0.05 and p < 0.02, respectively). CONCLUSIONS: These results indicate that divers with preexisting small lung cysts and/or end-expiratory flow limitation may be at risk of PBT.

Barotrauma↗

Glucoregulation and simulated diving in the harbor seal Phoca vitulina.

Plasma glucose, glucagon, and insulin concentrations were measured in the harbor seal, Phoca vitulina, during a 6-min dive and a 30-min recovery period. Studies were performed in the fasting state and following intravenous glucose. During diving in the fasting state, values of plasma glucose, glucagon, and insulin are not significantly different from prediving values, presumably because of the loss of perfusion to the pancreas. In the postdiving period, plasma glucagon increases significantly. The increased glucagon levels reach a peak at 6 min into the postdiving period. By 4 min postdiving, plasma glucose values increase. The hyperglycemia appears to be at least partially related to hyperglucagonemia. Despite hyperglycemia, insulin levels do not change significantly. The net effect is to increase glucose availability, particularly for non-insulin-requiring tissues (brain). Preadministration of glucose eliminates the postdiving increase in glucagon. Plasma glucose and insulin concentrations also show no significant changes during the diving and postdiving periods. These results suggest that hormonally mediated glucose conservation serves to maintain brain glucose supplies and to restore peripheral carbohydrate stores during the postdiving period.

Animals↗

Cardiovascular adjustments to laboratory diving in beavers and nutria.

Beavers (Castor canadensis) and nutria (Myocastor coypus) were anesthetized with halothane and catheters placed in the left ventricle, aorta and pulmonary artery, right ventricle or right atrium. The animals were strapped to a board and following recovery from anesthesia the following measurements were taken: regional distribution of blood flow, cardiac output, O2 consumption, arterial and venous blood gases, and pH. The animal was then immersed in 15-20 degrees C water for up to 2.75 min (nutria) or 4 min (beaver) and the measurements repeated. Heart rate and cardiac output decreased by 80 and 75%, respectively. Arterial and venous oxygen partial pressure and content fell as did pH whereas CO2 pressures rose during diving. Oxygen consumption at rest was 124 and 102% of that predicted on the basis of body mass for the beaver and nutria, respectively. Rate of decline of O2 stores during diving decreased by 93% in beavers and 89% in nutria compared to the predive value. Regional blood flow decreased to all organs except the adrenals, heart, and lungs. Blood flow to the brain increased during diving.

Adaptation, Physiological↗

Thermoregulatory control of expired air temperature in diving harp seals.

Expired air temperature (Tex), metabolic rate (MR), and skin (Ts) and body (Tb; rectal) temperatures were recorded in four or five young (1-2 yr) harp seals (Phoca groenlandica) in air [mean air temperature (Ta) = -30, -10, or 10 degrees C] and in water [mean water temperature (Tw) = 2.3 or 24.8 degrees C, with Ta = -30, -10, 0, or 10 degrees C]. Apparent lower critical temperature was below -10 degrees C in air. Above this Ta, mean MR was 5.85 W.kg-0.75 (2.23 W.kg-1), while mean MR was 12.56 W.kg-0.75 (4.69 W.kg-1) at Ta -30 degrees C. When seals were immersed in water of 2.3 degrees C, mean MR was 6.13 W.kg-0.75 (2.31 W.kg-1), regardless of Ta. At Ta -30, -10, and 10 degrees C, mean Tex in air were 9.5, 13.0, and 25.0 degrees C, respectively. The corresponding values for seals in water (Tw = 2.3 degrees C) were 8.0, 9.5, and 15.5 degrees C, respectively. The low Tex recorded at Ta -30 and -10 degrees C in air and at all Ta in water (Tw = 2.3 degrees C) suggests that heat was conserved by nasal heat exchange. At Ta 10 degrees C, mean Tex of seals in air was approximately 10 degrees C higher than mean Tex of seals in water (Tw = 2.3 degrees C). Furthermore, seals subjected to a Tw of 24.8 degrees C at Ta 0 degrees C had a mean Tex 10 degrees C higher than when subjected to Tw 2.3 degrees C at the same Ta. These observations suggest that Tex in seals is under thermoregulatory control. In a series of forced dives of up to 5-min duration Tex was found to be the same before and after the dive regardless of dive duration and Ta.

Animals↗

Effects of growth, diving history, and high altitude on blood oxygen capacity in harbor seals.

Blood volume and body composition measurements were carried out on diving and nondiving harbor seals, Phoca vitulina. There were small, statistically significant differences in several red blood cell parameters between diving and nondiving seals, but the effect was superimposed on uniformly large blood volumes ranging from 11 to 12% of body weight found in all the animals. It is suggested that the large blood oxygen transport and storage capacity is not so much a direct result of the apneic diving experience, but an expression of phylogenetic control. The development of blood volume during early growth was closely related to the increase in lean body mass. In spite of the inherently large blood volume, this marine mammal when exposed to high altitude for 85 days at an elevation of 3.1 km above sea level appeared to exhibit typical mammalian erythropoietic and ventilatory responses to long-term, mild hypoxia. Thus, it probably cannot be considered to be preacclimatized to high altitude.

Age Factors↗

Microcomputer monitor and blood sampler for free-diving Weddell seals.

The equipment used for the first sampling of arterial blood at depth on free-diving Weddell seals Leptonychotes weddelli is described. Blood was withdrawn through an aortic catheter by a submersible, peristaltic roller pump and stored in a single- or multiple-sample collection device. The multiple sampler allowed up to eight individual blood samples to be collected during a single dive. The blood pump was controlled by a dedicated microcomputer that allowed initiation of blood sampling at flexible combinations of depth and/or time during either the descending or ascending phase of the dive. The dedicated microcomputer also recorded swimming depth, velocity, heart rate, and body temperature at selectable time intervals. These data were transmitted to a laboratory computer, and blood samples were retrieved, when the seal surfaced to breathe.

Animals↗

Diving response and arterial oxygen saturation during apnea and exercise in breath-hold divers.

This study addressed the effects of apnea in air and apnea with face immersion in cold water (10 degrees C) on the diving response and arterial oxygen saturation during dynamic exercise. Eight trained breath-hold divers performed steady-state exercise on a cycle ergometer at 100 W. During exercise, each subject performed 30-s apneas in air and 30-s apneas with face immersion. The heart rate and arterial oxygen saturation decreased and blood pressure increased during the apneas. Compared with apneas in air, apneas with face immersion augmented the heart rate reduction from 21 to 33% (P < 0.001) and the blood pressure increase from 34 to 42% (P < 0.05). The reduction in arterial oxygen saturation from eupneic control was 6.8% during apneas in air and 5.2% during apneas with face immersion (P < 0.05). The results indicate that augmentation of the diving response slows down the depletion of the lung oxygen store, possibly associated with a larger reduction in peripheral venous oxygen stores and increased anaerobiosis. This mechanism delays the fall in alveolar and arterial PO(2) and, thereby, the development of hypoxia in vital organs. Accordingly, we conclude that the human diving response has an oxygen-conserving effect during exercise.

Adult↗

Blubber and buoyancy: monitoring the body condition of free-ranging seals using simple dive characteristics.

Elephant seals regularly perform dives during which they spend a large proportion of time drifting passively through the water column. The rate of vertical change in depth during these "drift" dives is largely a result of the proportion of lipid tissue in the body, with fatter seals having higher (more positive or less negative) drift rates compared with leaner seals. We examined the temporal changes in drift rates of 24 newly weaned southern elephant seal (Mirounga leonina) pups during their first trip to sea to determine if this easily recorded dive characteristic can be used to continuously monitor changes in body composition of seals throughout their foraging trips. All seals demonstrated a similar trend over time: drift rates were initially positive but decreased steadily over the first 30-50 days after departure (Phase 1), corresponding to seals becoming gradually less buoyant. Over the following approximately 100 days (Phase 2), drift rates again increased gradually, while during the last approximately 20-45 days (Phase 3) drift rates either remained constant or decreased slightly. The daily rate of change in drift rate was negatively related to the daily rate of horizontal displacement (daily travel rate), and daily travel rates of more than approximately 80 km were almost exclusively associated with negative changes in drift rate. We developed a mechanistic model based on body compositions and morphometrics measured in the field, published values for the density of seawater and various body components, and values of drag coefficients for objects of different shapes. We used this model to examine the theoretical relationships between drift rate and body composition and carried out a sensitivity analysis to quantify errors and biases caused by varying model parameters. While variations in seawater density and uncertainties in estimated body surface area and volume are unlikely to result in errors in estimated lipid content of more than +/-2.5%, variations in drag coefficient can lead to errors of >or =10%. Finally, we compared the lipid contents predicted by our model with the lipid contents measured using isotopically labelled water and found a strong positive correlation. The best-fitting model suggests that the drag coefficient of seals while drifting passively is between approximately 0.49 (roughly corresponding to a sphere-shaped object) and 0.69 (a prolate spheroid), and we were able to estimate relative lipid content to within approximately +/-2% lipid. Our results suggest that this simple method can be used to estimate the changes in lipid content of free-ranging seals while at sea and may help improve our understanding of the foraging strategies of these important marine predators.

Adipose Tissue↗

Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus.

Drag and buoyancy are two primary external forces acting on diving marine mammals. The strength of these forces modulates the energetic cost of movement and may influence swimming style (gait). Here we use a high-resolution digital tag to record depth, 3-D orientation, and sounds heard and produced by 23 deep-diving sperm whales in the Ligurian Sea and Gulf of Mexico. Periods of active thrusting versus gliding were identified through analysis of oscillations measured by a 3-axis accelerometer. Accelerations during 382 ascent glides of five whales (which made two or more steep ascents and for which we obtained a measurement of length) were strongly affected by depth and speed at Reynold's numbers of 1.4-2.8x10(7). The accelerations fit a model of drag, air buoyancy and tissue buoyancy forces with an r(2) of 99.1-99.8% for each whale. The model provided estimates (mean +/- S.D.) of the drag coefficient (0.00306+/-0.00015), air carried from the surface (26.4+/-3.9 l kg(-3) mass), and tissue density (1030+/-0.8 kg m(-3)) of these five animals. The model predicts strong positive buoyancy forces in the top 100 m of the water column, decreasing to near neutral buoyancy at 250-850 m. Mean descent speeds (1.45+/-0.19 m s(-1)) were slower than ascent speeds (1.63+/-0.22 m s(-1)), even though sperm whales stroked steadily (glides 5.3+/-6.3%) throughout descents and employed predominantly stroke-and-glide swimming (glides 37.7+/-16.4%) during ascents. Whales glided more during portions of dives when buoyancy aided their movement, and whales that glided more during ascent glided less during descent (and vice versa), supporting the hypothesis that buoyancy influences behavioural swimming decisions. One whale rested at approximately 10 m depth for more than 10 min without fluking, regulating its buoyancy by releasing air bubbles.

Acceleration↗

The effects of intense wing molt on diving in alcids and potential influences on the evolution of molt patterns.

Large and medium-sized alcids have a very intense wing molt wherein many flight feathers are shed in rapid succession and wing surface area is reduced by as much as 40%. Although these birds are rendered flightless during wing molt, they must still use their wings to propel themselves underwater. A molt-induced loss of wing area could simply reduce wing propulsion such that more muscular work would be required to maintain a given speed. Alternatively, molt could reduce drag on the wings, making a bird more penguin-like and actually enhancing diving ability. I addressed this issue by filming captive common guillemots Uria aalge and tufted puffins Fratercula cirrhata using an array of video cameras to plot the birds' movements in three dimensions. From these coordinate data I calculated swimming velocities, angles of descent and absolute depths. These values allowed me to estimate the forces due to drag and buoyancy that must be counteracted by flapping, which in turn yielded estimates of the amount of work generated during each flap as well as the average power and cost of transport. Within-bird comparisons of diving performance when wings were intact and during several stages of wing molt indicated that molt is associated with more frequent flapping, reduced displacement during the flap cycle, and possibly reduced work per flap. These negative effects on diving may explain why primary and secondary molts were offset in the birds I studied such that the period during which all of the flight feathers are effectively missing is minimized.

Analysis of Variance↗