Tracking of marine predators to protect Southern Ocean ecosystems.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change1,2. Mitigation requires the identification and protection of Areas of Ecological Significance (AESs), which have so far not been determined at the ocean-basin scale. Here, using assemblage-level tracking of marine predators, we identify AESs for this globally important region and assess current threats and protection levels. Integration of more than 4,000 tracks from 17 bird and mammal species reveals AESs around sub-Antarctic islands in the Atlantic and Indian Oceans and over the Antarctic continental shelf. Fishing pressure is disproportionately concentrated inside AESs, and climate change over the next century is predicted to impose pressure on these areas, particularly around the Antarctic continent. At present, 7.1% of the ocean south of 40°S is under formal protection, including 29% of the total AESs. The establishment and regular revision of networks of protection that encompass AESs are needed to provide long-term mitigation of growing pressures on Southern Ocean ecosystems.

Adaptations to deep and prolonged diving in phocid seals.

This Review focuses on the original papers that have made a difference to our thinking and were first in describing an adaptation to diving, and less on those that later repeated the findings with better equipment. It describes some important anatomical peculiarities of phocid seals, as well as their many physiological responses to diving. In so doing, it is argued that the persistent discussions on the relevance and differences between responses seen in forced dives in the laboratory and those during free diving in the wild are futile. In fact, both are two sides of the same coin, aimed at protecting the body against asphyxic insult and extending diving performance.

Waiting for the Sun: the circannual programme of reindeer is delayed by the recurrence of rhythmical melatonin secretion after the arctic night.

At temperate latitudes, the annual cycle of day length synchronizes circannual rhythms, and, in mammals, this is mediated via nocturnal production of the pineal hormone melatonin, proportional to the length of the night. Here, we studied circannual synchronization in an arctic species, the reindeer (Rangifer tarandus tarandus), which ceases to produce a rhythmic melatonin signal when it is exposed to extended periods of continuous midwinter darkness and continuous midsummer light. Using food intake, antler growth and moult as endpoints, we demonstrate that when animals living at 70°N are transferred from natural photoperiods in late autumn to either continuous light or continuous darkness, they undergo a conspicuous acceleration of the circannual programme. We conclude that rhythmical melatonin secretion, recommencing when the Sun reappears late in January, is required for proper timing of spring physiological responses, through a delaying effect on the circannual programme set in motion during the preceding autumn.

Ascending aorta of hooded seals with particular emphasis on its vasa vasorum.

The pressure-volume relationship in the ascending aorta ("windkessel") of the hooded seal was determined and the morphology of its vasa vasorum described in some detail. We found that the ascending aorta has a high compliance and can easily accommodate the entire stroke volume when the peripheral vascular resistance becomes much increased and maintain perfusion pressure during the much extended diastole and thereby reduce cardiac stroke work during diving. We also found that the 3- to 5-mm thick wall of the ascending aorta had a very elaborate vasa vasorum interna with a hitherto undescribed vascular structure that penetrates the entire vascular wall. If similar structures with similar importance for the nutrition of the wall of the vessel are found in humans, important implications for the understanding of pathological conditions, such as aneurisms, may be indicated.

Adaptations to polar life in mammals and birds.

This Review presents a broad overview of adaptations of truly Arctic and Antarctic mammals and birds to the challenges of polar life. The polar environment may be characterized by grisly cold, scarcity of food and darkness in winter, and lush conditions and continuous light in summer. Resident animals cope with these changes by behavioural, physical and physiological means. These include responses aimed at reducing exposure, such as 'balling up', huddling and shelter building; seasonal changes in insulation by fur, plumage and blubber; and circulatory adjustments aimed at preservation of core temperature, to which end the periphery and extremities are cooled to increase insulation. Newborn altricial animals have profound tolerance to hypothermia, but depend on parental care for warmth, whereas precocial mammals are well insulated and respond to cold with non-shivering thermogenesis in brown adipose tissue, and precocial birds shiver to produce heat. Most polar animals prepare themselves for shortness of food during winter by the deposition of large amounts of fat in times of plenty during autumn. These deposits are governed by a sliding set-point for body fatness throughout winter so that they last until the sun reappears in spring. Polar animals are, like most others, primarily active during the light part of the day, but when the sun never sets in summer and darkness prevails during winter, high-latitude animals become intermittently active around the clock, allowing opportunistic feeding at all times. The importance of understanding the needs of the individuals of a species to understand the responses of populations in times of climate change is emphasized.

On how whales avoid decompression sickness and why they sometimes strand.

Whales are unique in that the supply of blood to the brain is not by the internal carotid arteries, but by way of thoracic and intra-vertebral arterial retia. We found in the harbor porpoise (Phocoena phocoena) that these retia split up into smaller anastomosing vessels and thin-walled sinusoid structures that are embedded in fat. The solubility of nitrogen is at least six times larger in fat than in water, and we suggest that nitrogen in supersaturated blood will be absorbed in the fat, by diffusion, during the very slow passage of the blood through the arterial retia. Formation of nitrogen bubbles that may reach the brain is thereby avoided. We also suggest that mass stranding of whales may be due to disturbances to their normal dive profiles, resulting in extra release of nitrogen that may overburden the nitrogen 'trap' and allow bubbles to reach the brain and cause abnormal behavior.

Experimental reduction of food quality is not compensated with increased food intake in high-Arctic muskoxen (Ovibos moschatus).

A total of four barren adult female muskoxen (Ovibos moschatus) were used over a period of 2 years for the purpose of the present study. During the first year, the natural changes in appetite (ad libitum intake of standard pelleted reindeer feed) and body mass were determined in two of the animals. During the second year, the effect of reduced food quality on ad libitum food intake was tested in all four animals in July when the appetite had been found to be at a high. We found that the experimentally reduced food quality was not compensated with increased food intake in these large high-Arctic herbivores.

The venous system of seals, with new ideas on the significance of the extradural intravertebral vein.

The structural features of the venous system of seals, including such specialties as a caval sphincter, a huge posterior caval vein and hepatic sinuses, venous plexuses and a huge extradural intravertebral vein, are described and functional aspects of these features in relation to diving habits are discussed. A number of old concepts are discarded and new ones are proposed, particularly with regard to the significance of the extradural intravertebral vein, which, contrary to common current belief, probably is of little importance during diving.

Regulation of brain temperature in winter-acclimatized reindeer under heat stress.

Reindeer (Rangifer tarandus) are protected against the Arctic winter cold by thick fur of prime insulating capacity and hence have few avenues of heat loss during work. We have investigated how these animals regulate brain temperature under heavy heat loads. Animals were instrumented for measurements of blood flow, tissue temperatures and respiratory frequency (f) under full anaesthesia, whereas measurements were also made in fully conscious animals while in a climatic chamber or running on a treadmill. At rest, brain temperature (T(brain)) rose from 38.5±0.1°C at 10°C to 39.5±0.2°C at 50°C, while f increased from ×7 to ×250 breaths min(-1), with a change to open-mouth panting (OMP) at T(brain) 39.0±0.1°C, and carotid and sublingual arterial flows increased by 160% and 500%, respectively. OMP caused jugular venous and carotid arterial temperatures to drop, presumably owing to a much increased respiratory evaporative heat loss. Angular oculi vein (AOV) flow was negligible until T(brain) reached 38.9±0.1°C, but it increased to 0.81 ml min(-1) kg(-1) at T(brain) 39.2±0.2°C. Bilateral occlusion of both AOVs induced OMP and a rise in T(brain) and f at T(brain) >38.8°C. We propose that reindeer regulate body and, particularly, brain temperature under heavy heat loads by a combination of panting, at first through the nose, but later, when the heat load and the minute volume requirements increase due to exercise, primarily through the mouth and that they eventually resort to selective brain cooling.

Circadian organization in reindeer.

The light/dark cycle of day and night synchronizes an internal 'biological clock' that governs daily rhythms in behaviour, but this form of regulation is denied to polar animals for most of the year. Here we demonstrate that the continuous lighting conditions of summer and of winter at high latitudes cause a loss in daily rhythmic activity in reindeer living far above the Arctic Circle. This seasonal absence of circadian rhythmicity may be a ubiquitous trait among resident polar vertebrates.

Selective brain cooling and its vascular basis in diving seals.

Brain (T(brain)), intra-aorta (T(aorta)), latissimus dorsi muscle (T(m)) and rectal temperature (T(r)) were measured in harp (Pagophilus groenlandicus) and hooded (Cystophora cristata) seals during experimental dives in 4 degrees C water. The median brain cooling was about 1 degrees C during 15 min diving, but in some cases it was as much as 2.5 degrees C. Cooling rates were slow for the first couple of minutes, but increased significantly after about 5 min of diving. The onset of cooling sometimes occurred before the start of the dive, confirming that the cooling is under cortical control, like the rest of the diving responses. T(aorta) also fell significantly, and was always lower than T(brain), while T(m) was fairly stable during dives. Detailed studies of the vascular anatomy of front flippers revealed that brachial arterial blood can be routed either through flipper skin capillaries for nutritive purposes and return through sophisticated vascular heat exchangers to avoid heat loss to the environment, or, alternatively, through numerous arterio-venous shunts in the skin and return by way of large superficial veins, which then carry cold blood to the heart. In the latter situation the extent to which the brain is cooled is determined by the ratio of carotid to brachial arterial blood flow, and water temperature, and the cooling is selective in that only those organs that are circulated will be cooled. It is concluded that T(brain) is actively down-regulated during diving, sometimes by as much as 2.5 degrees C, whereby cerebral oxygen requirements may be reduced by as much as 25% during extended dives.

Microbial degradation of usnic acid in the reindeer rumen.

Reindeer (Rangifer tarandus) eat and utilize lichens as an important source of energy and nutrients in winter. Lichens synthesize and accumulate a wide variety of phenolic secondary compounds, such as usnic acid, as a defense against herbivores and to protect against damage by UV-light in solar radiation. We have examined where and to what extent these phenolic compounds are degraded in the digestive tract of the reindeer, with particular focus on usnic acid. Three male reindeer were given ad libitum access to a control diet containing no usnic acid for three weeks and then fed lichens ad libitum (primarily Cladonia stellaris) containing 9.1 mg/g DM usnic acid for 4 weeks. Usnic acid intake in reindeer on the lichen diet was 91-117 mg/kg BM/day. In spite of this, no trace of usnic acid or conjugates of usnic acid was found either in fresh rumen fluid, urine, or feces. This suggests that usnic acid is rapidly degraded by rumen microbes, and that it consequently is not absorbed by the animal. This apparent ability to detoxify lichen phenolic compounds may gain increased importance with future enhanced UV-B radiation expected to cause increased protective usnic acid/phenol production in lichens.

A possible cause of Sudden Infant Death Syndrome.

It is suggested that an orienting response to loud sound causes apnea, which, in already asphyxic infants, triggers a maximal secondary chemoreceptor response, with massive vagal stimulation of the heart, which causes heart arrest.

The retrospective analysis of Antarctic tracking data project.

The Retrospective Analysis of Antarctic Tracking Data (RAATD) is a Scientific Committee for Antarctic Research project led jointly by the Expert Groups on Birds and Marine Mammals and Antarctic Biodiversity Informatics, and endorsed by the Commission for the Conservation of Antarctic Marine Living Resources. RAATD consolidated tracking data for multiple species of Antarctic meso- and top-predators to identify Areas of Ecological Significance. These datasets and accompanying syntheses provide a greater understanding of fundamental ecosystem processes in the Southern Ocean, support modelling of predator distributions under future climate scenarios and create inputs that can be incorporated into decision making processes by management authorities. In this data paper, we present the compiled tracking data from research groups that have worked in the Antarctic since the 1990s. The data are publicly available through biodiversity.aq and the Ocean Biogeographic Information System. The archive includes tracking data from over 70 contributors across 12 national Antarctic programs, and includes data from 17 predator species, 4060 individual animals, and over 2.9 million observed locations.

Immobilization of muskoxen (Ovibos moschatus) with etorphine and xylazine.

One hundred and thirty three "wild" muskoxen, 81 of which of known body mass, were successfully immobilized using etorphine (M99), and xylazine (Rompun®), delivered by use of a dart gun. A dose of 0.05 mg/kg M99, supplemented by 0.15 mg/kg Rompun was found to be very effective. This dose is much higher than currently recommended e.g. by Handbook of Wildlife Chemical Immobilization.

Inhibition of shivering in hypothermic seals during diving.

The mammalian response to hypothermia is increased metabolic heat production, usually by way of muscular activity, such as shivering. Seals, however, have been reported to respond to diving with hypothermia, which in other mammals under other circumstances would have elicited vigorous shivering. In the diving situation, shivering could be counterproductive, because it obviously would increase oxygen consumption and therefore reduce diving capacity. We have measured the electromyographic (EMG) activity of three different muscles and the rectal and brain temperature of hooded seals (Cystophora cristata) while they were exposed to low ambient temperatures in a climatic chamber and while they performed a series of experimental dives in cold water. In air, the seals had a normal mammalian shivering response to cold. Muscles were recruited in a sequential manner until body temperature stopped dropping. Shivering was initiated when rectal temperature fell below 35.3 +/- 0.6 degrees C (n = 6). In the hypothermic diving seal, however, the EMG activity in all of the muscles that had been shivering vigorously before submergence was much reduced, or stopped altogether, whereas it increased again upon emergence but was again reduced if diving was repeated. We conclude that shivering is inhibited during diving to allow a decrease in body temperature whereby oxygen consumption is decreased and diving capacity is extended.