The physiology of underwater diving is the physiological adaptations to diving of air-breathing vertebrates that have returned to the ocean from terrestrial lineages. They are a diverse group that include sea snakes, sea turtles, the marine iguana, saltwater crocodiles, penguins, pinnipeds, cetaceans, sea otters, manatees and dugongs. All known diving vertebrates dive to feed, and the extent of the diving in terms of depth and duration are influenced by feeding strategies, but also, in some cases, with predator avoidance. Diving behaviour is inextricably linked with the physiological adaptations for diving and often the behaviour leads to an investigation of the physiology that makes the behaviour possible, so they are considered together where possible. Most diving vertebrates make relatively short shallow dives. Sea snakes, crocodiles, and marine iguanas only dive in inshore waters and seldom dive deeper than 10 meters (33 feet). Some of these groups can make much deeper and longer dives. Emperor penguins regularly dive to depths of 400 to 500 meters (1,300 to 1,600 feet) for 4 to 5 minutes, often dive for 8 to 12 minutes, and have a maximum endurance of about 22 minutes. Elephant seals stay at sea for between 2 and 8 months and dive continuously, spending 90% of their time underwater and averaging 20 minutes per dive with less than 3 minutes at the surface between dives. Their maximum dive duration is about 2 hours and they routinely feed at depths between 300 and 600 meters (980 and 1,970 feet), though they can exceed depths of 1,600 meters (5,200 feet). Beaked whales have been found to routinely dive to forage at depths between 835 and 1,070 meters (2,740 and 3,510 feet), and remain submerged for about 50 minutes. Their maximum recorded depth is 1,888 meters (6,194 feet), and the maximum duration is 85 minutes. Air-breathing marine vertebrates that dive to feed must deal with the effects of pressure at depth, hypoxia during apnea, and the need to find and capture their food. Adaptations to diving can be associated with these three requirements. Adaptations to pressure must deal with the mechanical effects of pressure on gas-filled cavities, solubility changes of gases under pressure, and possible direct effects of pressure on the metabolism, while adaptations to breath-hold capacity include modifications to metabolism, perfusion, carbon dioxide tolerance, and oxygen storage capacity. Adaptations to find and capture food vary depending on the food, but deep-diving generally involves operating in a dark environment. Diving vertebrates have increased the amount of oxygen stored in their internal tissues. This oxygen store has three components; oxygen contained in the air in the lungs, oxygen stored by haemoglobin in the blood, and by myoglobin, in muscle tissue, The muscle and blood of diving vertebrates have greater concentrations of haemoglobin and myoglobin than terrestrial animals. Myoglobin concentration in locomotor muscles of diving vertebrates is up to 30 times more than in terrestrial relatives. Haemoglobin is increased by both a relatively larger amount of blood and a larger proportion of red blood cells in the blood compared with terrestrial animals. The highest values are found in the mammals which dive deepest and longest. Body size is a factor in diving ability. A larger body mass correlates to a relatively lower metabolic rate, while oxygen storage is directly proportional to body mass, so larger animals should be able to dive for longer, all other things being equal. Swimming efficiency also affects diving ability, as low drag and high propulsive efficiency requires less energy for the same dive. Burst and glide locomotion is also often used to minimise energy consumption, and may involve using positive or negative buoyancy to power part of the ascent or descent. The responses seen in seals diving freely at sea are physiologically the same as those seen during forced dives in the laboratory. They are not specific to immersion in water, but are protective mechanisms against asphyxia which are common to all mammals but more effective and developed in seals. The extent to which these responses are expressed depends greatly on the seal's anticipation of dive duration. The regulation of bradycardia and vasoconstriction of the dive response in both mammals and diving ducks can be triggered by facial immersion, wetting of the nostrils and glottis, or stimulation of trigeminal and glossopharyngeal nerves. Animals cannot convert fats to glucose, and in many diving animals, carbohydrates are not readily available from the diet, nor stored in large quantities, so as they are essential for anaerobic metabolism, they could be a limiting factor. Decompression sickness (DCS) is a disease associated with metabolically inert gas uptake at pressure, and its subsequent release into the tissues in the form of bubbles. Marine mammals were thought to be relatively immune to DCS due to anatomical, physiological and behavioural adaptations that reduce tissue loading with dissolved nitrogen during dives, but observations show that gas bubbles may form, and tissue injury may occur under certain circumstances. Decompression modelelling using measured dive profiles predict the possibility of high blood and tissue nitrogen tensions.
Range of performance Strong allometric relationships have been observed between body mass and maximum diving depths and durations in marine mammals, birds and marine turtles. Exceptions can be seen in baleen whales, eared seals and flighted diving birds, which showed no correlation between size and maximum diving depth. Penguins showed the highest correlations of diving capacity with body mass, followed by toothed whales and true seals. Baleen whales show a correlation between mass and maximum dive duration. Comparisons between the groups show that alcids, penguins, and true seals are exceptional divers relative to their masses and that baleen whales dive to shallower depths and for shorter durations than would be expected from their size. Many of these variations can be explained by adaptation to ecological feeding niches exploited by the animals.
Marine mammals
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