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Fur seals may look like they’re simply resting after exhausting hunting trips at sea, but their bodies are secretly working overtime。 Scientists discovered that hours after returning to land, the seals’ heart rates suddenly surge — sometimes doubling — as they recover from the intense physical stress of deep diving。 The findings suggest that seals

Abstract: Scuba diving on coral reefs is an increasingly lucrative element of tourism in the tropics, but divers can damage the reefs on which tourism depends. By studying the effects of diving we can determine what level of use is justifiable in balancing objectives of economic gain and conservation. Off the Caribbean island of Bonaire we compared coral and fish communities between undived reserves and environmentally similar dive sites where maximum use reached 6000 dives per site per year. At these levels of diving, direct physical damage to reefs was relatively minor. There were more loose fragments of living coral in dive sites than reserves and more abraded coral in high‐ than low‐use areas. Diving had no significant effect on reef fish communities. Between 1991 and 1994, diving intensity increased 70% and coral cover declined in two of three dive sites and in all three reserves, suggesting a background stress unrelated to tourism. There was a significant decline in the proportion of old colonies of massive coral species within dive sites (19.2% loss), compared to a smaller loss in reserves (6.7%). Branching corals increased by 8.2% in dive sites, compared with 2.2% in reserves. Despite close management of reefs, diving is changing the character of Bonaire's reefs by allowing branching corals to increase at the expense of large, massive colonies. The impact of background stresses on massive corals seems to have been greater in the presence of diving. Other studies have linked disease infection to coral tissue damage, and the higher rates of abrasion we recorded in dived sites could have rendered corals there more susceptible to disease, thus mediating the decline of massive corals. Our study shows that even relatively low levels of diving can have pronounced effects manifested in shifts in dominance patterns rather than loss of overall coral cover. Bonaire's reefs have among the highest coral cover and greatest representation of ancient coral colonies of reefs anywhere in the Caribbean. Conserving the character of these reefs may require tighter controls on diving intensity.

The sections in this article are: 1 Historical Landmarks 2 The Basic Problem for Diving Animals 3 Decreased Sensitivity to Asphyxia 4 Oxygen Stores and Principles of Their Utilization 5 Cardiovascular Adjustments During and After Diving 5.1 General Aspects 5.2 Cardiac Output 5.2.1 General Considerations 5.2.2 Cardiac Output During Diving 5.2.3 Cardiac Output in Recovery 5.3 Regional Circulation During Diving and on Emersion 5.3.1 General Aspects 5.3.2 Cerebral Circulation 5.3.3 Myocardial Blood Supply 5.3.4 Renal Blood Flow 5.3.5 Gastrointestinal and Liver Blood Flow 5.3.6 Skeletal Muscle Circulation 5.3.7 Cutaneous Circulation 5.3.8 Adrenal Circulation 5.3.9 Uterine-Placental Circulation 5.3.10 Vascular Changes on Emersion 6 Morphological Specializations in Vascular Beds of Diving Animals 6.1 Arteries 6.2 Shunts 6.3 Veins 7 Metabolic Consequences 8 Central Integration of Cardiovascular Responses 8.1 Medullary Reflex Mechanisms 8.1.1 General Aspects 8.1.2 Initial Diving Responses 8.1.3 Chemoreceptor Reinforcement of Initial Diving Responses 8.2 Suprabulbar Influences 8.3 Other Reflex Influences 8.3.1 Cardiac Volume Receptors 8.3.2 Arterial Baroreceptors 8.3.3 Pulmonary-Thoracic Stretch Receptors 9 Diving Responses in Humans and Some Clinical Applications 9.1 Responses in Humans 9.2 Clinical Applications

Maximum diving depths and durations were examined in relation to body mass for birds, marine mammals, and marine turtles. There were strong allometric relationships between these parameters (log 10 transformed) among air-breathing vertebrates (r = 0.71, n = 111 for depth; r = 0.84, n = 121 for duration), although there was considerable scatter around the regression lines. Many of the smaller taxonomic groups also had a strong allometric relationship between diving capacity (maximum depth and duration) and body mass. Notable exceptions were mysticete cetaceans and diving/flying birds, which displayed no relationship between maximum diving depth and body mass, and otariid seals, which showed no relationship between maximum diving depth or duration and body mass. Within the diving/flying bird group, only alcids showed a significant relationship (r = 0.81, n = 9 for depth). The diving capacities of penguins had the highest correlations with body mass (r = 0.81, n = 11 for depth; r = 0.93, n = 9 for duration), followed by those of odontocete cetaceans (r = 0.75, n = 21 for depth; r = 0.84, n = 22 for duration) and phocid seals (r = 0.70, n = 15 for depth; r = 0.59, n = 16 for duration). Mysticete cetaceans showed a strong relationship between maximum duration and body mass (r = 0.84, n = 9). Comparisons across the various groups indicated that alcids, penguins, and phocids are all exceptional divers relative to their masses and that mysticete cetaceans dive to shallower depths and for shorter periods than would be predicted from their size. Differences among groups, as well as the lack of relationships within some groups, could often be explained by factors such as the various ecological feeding niches these groups exploit, or by variations in the methods used to record their behavior.

The leading textbook of diving medicine, by international experts, has been completely revised and updated. It provides a comprehensive account relating the basic medical sciences to clinical conditions associated with diving. In-depth coverage of the physiological basis for safe diving, the pathophysiological basis for diving illnesses and the management of diving accidents is included. Features new chapters on fitness to dive, long term health effects of diving, and management of diving accidents.

This review concentrates on the physiological responses, and their control, in freely diving birds and mammals that enable them to remain submerged and sometimes quite active for extended periods of time. Recent developments in technology have provided much detailed information on the behavior of these fascinating animals. Unfortunately, the advances in technology have been insufficient to enable physiologists to obtain anything like the same level of detail on the metabolic rate and physiological adjustments that occur during natural diving. This has led to much speculation and calculations based on many assumptions concerning usable oxygen stores and metabolic rate during diving, in an attempt to explain the observed behavior. Despite their shortcomings, these calculations have provided useful insights into the degree of adaptations of various species of aquatic birds and mammals. Many of them, e.g., ducks, smaller penguins, fur seals, and Weddell seals, seem able to metabolize aerobically, when diving, at approximately the same (if not greater) rate as they do at the surface. Their enhanced oxygen stores are able to support aerobic metabolism, at what would not be considered unusually low levels, for the duration of the dives, although there are probably circulatory readjustments to ensure that the oxygen stores are managed judiciously. For other species, such as the larger penguins, South Georgian shag, and female elephant seals, there is a general consensus that they must either be reducing their aerobic metabolic rate when diving, possibly by way of regional hypothermia, and/or producing ATP, at least partly, by anaerobiosis and metabolizing the lactic acid when at the surface (although this is hardly likely in the case of the female elephant seals). Circulation is the proximate regulator of metabolism during aerobic diving, and heart rate is the best single indicator of circulatory adjustment. During voluntary dives, heart rates range from extreme bradycardia to well above resting, reflecting metabolic performance. Efferent cardiac control is largely parasympathetic. Reflex cardiorespiratory responses are modulated by conditioning and habituation, but reflexes predominate during extended dives and during recovery, when gas exchange is maximized.

The purpose of this review is to outline the physiological responses associated with the diving response, its functional significance, and its cardiorespiratory control. This review is separated into four major sections. Section one outlines the diving response and its physiology. Section two provides support for the hypothesis that the primary role of the diving response is the conservation of oxygen. The third section describes how the diving response is controlled and provides a model that illustrates the cardiorespiratory interaction. Finally, the fourth section illustrates potential adaptations that result after regular exposure to an asphyxic environment. The cardiovascular and endocrine responses associated with the diving response and apnea are bradycardia, vasoconstriction, and an increase in secretion of suprarenal catecholamines. These responses require the integration of both the cardiovascular system and the respiratory system. The primary role of the diving response is likely to conserve oxygen for sensitive brain and heart tissue and to lengthen the time before the onset of serious hypoxic damage. We suggest that future research should be focused towards understanding the role of altered ventilatory responses in human breath-hold athletes as well as in patients suffering from sleep-disordered breathing.

The metabolic rates of freely diving Weddell seals were measured using modern methods of on-line computer analysis coupled to oxygen consumption instrumentation. Oxygen consumption values were collected during sleep, resting periods while awake and during diving periods with the seals breathing at the surface of the water in an experimental sea-ice hole in Antarctica. Oxygen consumption during diving was not elevated over resting values but was statistically about 1.5 times greater than sleeping values. The metabolic rate of diving declined with increasing dive duration, but there was no significant difference between resting rates and rates in dives lasting up to 82 min. Swimming speed, measured with a microprocessor velocity recorder, was constant in each animal. Calculations of the aerobic dive limit of these seals were made from the oxygen consumption values and demonstrated that most dives were within this theoretical limit. The results indicate that the cost of diving is remarkably low in Weddell seals relative to other diving mammals and birds.

The oxygen store/usage hypothesis suggests that larger animals are able to dive for longer and hence deeper because oxygen storage scales isometrically with body mass, whereas oxygen usage scales allometrically with an exponent <1 (typically 0.67-0.75). Previous tests of the allometry of diving tend to reject this hypothesis, but they are based on restricted data sets or invalid statistical analyses (which assume that every species provides independent information). Here we apply information-theoretic statistical methods that are phylogenetically informed to a large data set on diving variables for birds and mammals to describe the allometry of diving. Body mass is strongly related to all dive variables except dive:pause ratio. We demonstrate that many diving variables covary strongly with body mass and that they have allometric exponents close to 0.33. Thus, our results fail to falsify the oxygen store/usage hypothesis. The allometric relationships for most diving variables are statistically indistinguishable for birds and mammals, but birds tend to dive deeper than mammals of equivalent mass. The allometric relationships for all diving variables except mean dive duration are also statistically indistinguishable for all major taxonomic groups of divers within birds and mammals, with the exception of the procellariiforms, which, strictly speaking, are not true divers.

Human cold adaptation was studied by comparing maximal body insulation [ I = (rectal temp. – skin temp.) /rate of skin heat loss] of Korean diving women to insulation of Korean nondiving men and women and American men and women. Appropriate measurements were made during immersion in a constant-temperature bath cool enough to induce maximal cutaneous vasoconstriction without shivering. Subcutaneous fat was estimated from measurements of skin-fold thickness. Within each racial group there is a significant regression of I on fat thickness. Koreans had a significantly greater I than Americans of comparable fat thickness. Korean diving women had the same I as nondivers of comparable fat thickness. Korean women had significantly greater I than Korean men due, we believe, to thicker subcutaneous fat. This may be the reason why women and not men engage in diving. The only evidence for cold adaptation among diving women was their ability to withstand colder water immersion without shivering. Submitted on March 22, 1962

King Penguins are the second largest of all diving birds and share with their congener, Emperor Penguins, breeding habits strikingly different from other penguins. Our purpose was to determine the feeding behavior, energetics of foraging and the prey species, and compare these to other sympatric species of subantarctic divers. We determined: (1) general features of foraging behavior using time—depth recorders, velocity meters, and radio transmitters, (2) energetics by doubly labeled water, (3) food habits and energy content from stomach lavage samples, and (4) resting and swimming metabolic rate by oxygen consumption measurements. The average foraging cycle was ≈6 d, during which the mass gain of 30 birds was ≈2 kg. When at sea, the birds exhibit a marked pattern of shallow dives during the night, whereas deep dives of &gt;100 m only occurred during the day. Maximum depth measured from 34 birds and 18 537 dives was 304 m, and maximum dive duration from 23 birds and 11 874 dives was 7.7 min. The frequency distribution of dive depth was bimodal, with few dives between 40 and 100 m. Overall, swim velocities when a bird was at sea averaged 2.1 m/s (N = 5), while descent and ascent rates of change in depth averaged 0.6 m/s for dives &lt;60 m (N = 74) and 1.4 m/s for dives &gt;150 m (N = 90). Night feeding dives occurred at a rate of ≈20 dives/h, and deep dives occurred at a rate of ≈5 dives/h. The energy consumption rate while resting ashore was 3.3 W/kg (N = 3) or 1.6 times the predicted standard metabolic rate (SMR). The average energy consumption rate while away from the colony was 10 W/kg (N = 8) or 4.6 x SMR, compared to 4.3 x SMR estimated from a time—energy budget. The latter value is based on an average metabolic rate of 4.2 W/kg for three birds while resting in 5°C water and 9.6 W/kg while swimming at 2 m/s, which was extrapolated from the average of three birds swimming at 1 m/s. The average energy intake based on 9 stomach content samples was nearly 24.6 kJ/g dry mass. The main prey by number are myctophid fish of the species Krefftichthys anderssoni and Electrona carlsbergi. It was concluded that: (1) feeding begins ≈28 km from the colony, (2) prey is pursued night and day through its vertical movements, (3) vertical distribution of the prey is reflected closely by diving habits of the birds, (4) deep—diving, for unknown reasons, is an important component of foraging success, (5) diving capacities of King Penguins are remarkable compared to other birds and many pinnipeds, and (6) calculated foraging energetics can be closely estimated from time—energy budgets.

Changes in regional blood flow during simulated normobaric diving were studied in the conscious Antarctic Weddell seal (Leptonychotes weddelli) by injecting 25-microns radioactive microspheres into the left ventricle. Injections were performed before and 8--12 min after submersion of the head in iced seawater. Diving was associated with a fall in cardiac output from a mean control value of 39.8 +/- 10.2 to 5.6 +/- 3.4 l/min (mean +/- SD) and in heart rate from 52 +/- 15 to 15 +/- 4 beats/min. Blood flow to the splanchnic and peripheral vascular bed was reduced by more than 90%, cerebral blood flow was unchanged, right and left ventricular blood flow decreased by 85%, and adrenal blood flow decreased by 39%. The pulmonary fraction of the injected microsphere dose increased from 7.9 to 29.9% during diving. This may signify a large increase of peripheral arteriovenous shunting during the dive and/or increased bronchial artery blood flow. It is concluded that blood flow during diving is directed to organs and tissues according to their metabolic requirements.

Considered an essential resource by many in the field, Diving and Subaquatic Medicine remains the leading text on diving medicine, written to fulfil the requirements of any general physician wishing to advise their patients appropriately when a diving trip is planned, for those accompanying diving expeditions or when a doctor is required to assess

Understanding the physiology and genetics of human hypoxia tolerance has important medical implications, but this phenomenon has thus far only been investigated in high-altitude human populations. Another system, yet to be explored, is humans who engage in breath-hold diving. The indigenous Bajau people ("Sea Nomads") of Southeast Asia live a subsistence lifestyle based on breath-hold diving and are renowned for their extraordinary breath-holding abilities. However, it is unknown whether this has a genetic basis. Using a comparative genomic study, we show that natural selection on genetic variants in the PDE10A gene have increased spleen size in the Bajau, providing them with a larger reservoir of oxygenated red blood cells. We also find evidence of strong selection specific to the Bajau on BDKRB2, a gene affecting the human diving reflex. Thus, the Bajau, and possibly other diving populations, provide a new opportunity to study human adaptation to hypoxia tolerance. VIDEO ABSTRACT.