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Development of the blood and muscle oxygen stores in gray seals (Halichoerus grypus): implications for juvenile diving capacity and the necessity of a terrestrial postweaning fast.

To successfully transition from nursing to foraging, phocid seal pups must develop adequate diving physiology within the limited time between birth and their first independent foraging trip to sea. We studied the postpartum development of oxygen stores in gray seals (Halichoerus grypus, n=40) to better understand the ontogeny of diving capacity in phocids. Hemoglobin (Hb), hematocrit (Hct), blood volume (BV), and myoglobin (Mb) levels in newborn (3 d postpartum [DPP]) and newly weaned (17+/-0.4 DPP) pups were among the lowest measured across age classes. During the pups' terrestrial postweaning fast (PWF), Hb, Hct, mass-specific BV, and Mb increased by 28%, 21%, 13%, and 29%, respectively, resulting in a 35% increase in total body mass-specific oxygen stores and a 23% increase in calculated aerobic dive limit (CADL). Although Hb and Hct levels at the end of the PWF were nearly identical to those of yearlings, total body mass-specific oxygen stores and CADL of weaned pups departing for sea were only 66%-67% and 32%-62%, respectively, of those for yearlings and adult females. The PWF represents an integral component of the physiological development of diving capacity in phocids; however, newly independent phocids still appear to have limited diving capabilities at the onset of foraging.

Analysis of Variance↗

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice.

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1+/-0.2 degrees C (n=101 dives in five birds), pectoral muscle: 37.8+/-0.1 degrees C (n=71 dives in three birds) and axillary/brachial veins: 37.9+/-0.1 degrees C (n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=-0.59, P<0.05; range <1 degrees C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6+/-0.7 degrees C (n=157 dives in three birds), 20.2+/-1.2 degrees C (n=69 in three birds) and 35.2+/-0.2 degrees C (n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r=-0.29 to -0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r=-0.49 and -0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 degrees C during rest periods to a grand mean of 15.0+/-0.7 degrees C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r=-0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degrees C more closely than the anterior abdominal temperatures (19-30 degrees C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.

Animals↗

Field physiology: physiological insights from animals in nature.

Whereas comparative physiology documents the range of physiological variation across a range of organisms, field physiology provides insight into the actual mechanisms an organism employs to maintain homeostasis in its everyday life. This requires an understanding of an organism's natural history and is prerequisite to developing hypotheses about physiological mechanisms. This review focuses on a few areas of field physiology that exemplify how the underlying physiology could not have been understood without appropriate field measurements. The examples we have chosen highlight the methods and inference afforded by an application of this physiological analysis to organismal function in nature, often in extreme environments. The specific areas examined are diving physiology, the thermal physiology of large endothermic fishes, reproductive physiology of air breathing vertebrates, and endocrine physiology of reproductive homeostasis. These areas form a bridge from physiological ecology to evolutionary ecology. All our examples revolve around the central issue of physiological limits as they apply to organismal homeostasis. We view this theme as the cornerstone of physiological analysis and supply a number of paradigms on homeostasis that have been tested in the context of field physiology.

Animals↗

Instrumentation for the remote monitoring of physiological and behavioral variables.

Few commercial products are available for investigators who wish to monitor multiple physiological and behavioral variables in unrestrained subjects. When telemetry is not practical, e.g., in studies of at-sea diving physiology, one of the only options is to design and build a custom data-logging instrument. This paper describes how a data logger was developed for the successful long-term monitoring of dive depth, swim speed, heart rate, water temperature, and multiple body temperatures from free-ranging northern elephant seals. The task was facilitated by using a commercially available single-board computer designed specifically for portable multichannel data acquisition and, where possible, off-the-shelf sensors/transducers available with integrated signal-conditioning circuits. A smaller data logger for monitoring the electrocardiogram, body temperature, and dive behavior of double-crested cormorants is also described to illustrate the flexibility and simplicity of this approach. Although it is customized for diving animals, with incorporation of the appropriate sensors the basic design should be applicable to studies of comparative, environmental, or exercise physiology involving most medium-to-large animals, including humans.

Animals↗

A diving fatality due to oxygen toxicity during a "technical" dive.

An experienced diver drowned after a generalised seizure caused by oxygen toxicity during a 19-minute "technical" dive to a depth of 47 m. He had used a 50% oxygen/nitrogen gas mixture inappropriately during the dive. This case attests the risk of oxygen toxicity from oxygen-enriched air during deep dives, the shortcomings of the diver's equipment, and the need to examine, with knowledge of diving physiology and practice, both the body and equipment.

Diving↗

Convective oxygen transport and tissue oxygen consumption in Weddell seals during aerobic dives.

Unlike their terrestrial counterparts, marine mammals stop breathing and reduce their convective oxygen transport while performing activities (e.g. foraging, courtship, aggressive interactions, predator avoidance and migration) that require sustained power output during submergence. Since most voluntary dives are believed to remain aerobic, the goal of this study was to examine the potential importance of the dive response in optimizing the use of blood and muscle oxygen stores during dives involving different levels of muscular exertion. To accomplish this, we designed a numerical model based on Fick's principle that integrated cardiac output (Vb), regional blood flow, convective oxygen transport (Q(O2)), muscle oxymyoglobin desaturation and regional rates of oxygen consumption (VO2). The model quantified how the optimal matching or mismatching of QO2 to VO2 affected the aerobic dive limit (ADL). We chose an adult Weddell seal Leptonycotes weddellii on which to base our model because of available data on the diving physiology and metabolism of this species. The results show that the use of blood and muscle oxygen stores must be completed at the same time to maximize the ADL for each level of VO2. This is achieved by adjusting Vb (range 19-94 % of resting levels) and muscle QO2 according to the rate of muscle oxygen consumption (VMO2). At higher values of VMO2, Vb and muscle perfusion must increase to maintain an appropriate QO2/VO2 ratio so that available blood and muscle oxygen stores are depleted at the same time. Although the dive response does not sequester blood oxygen exclusively for brain and heart metabolism during aerobic dives, as it does during forced submersion, a reduction in Vb and muscle perfusion below resting levels is necessary to maximize the ADL over the range of diving VO2 (approximately 2-9 ml O2 min-1 kg-1). Despite the reduction in Vb, convective oxygen transport is adequate to maintain aerobic metabolism and normal function in the splanchnic organs, kidneys and other peripheral tissues. As a result, physiological homeostasis is maintained throughout the dive. The model shows that the cardiovascular adjustments known as the dive response enable the diving seal to balance the conflicting metabolic demands of (1) optimizing the distribution and use of blood and muscle oxygen stores to maximize the ADL over the normal range of diving VO2 and (2) ensuring that active muscle receives adequate oxygen as VMO2 increases.

Aerobiosis↗

[Current problems in prophylaxis and treatment of acute decompression disease of mild degree].

The authors discuss the causes leading to the rise of acute decompression illness in the amateur skin-divers. They include the insufficient training of most skin-drivers in the basis of diving physiology and medicine, the underwater dipping without consideration of individual sensitivity to decompression illness and nitrogen narcotic action, disorders in decompression regimens, non-observance of behavior rules after dipping, use of air transport immediately after dipping. The case of delayed treatment of decompression illness is described. The data concerning the possibility of chronic decompression illness formation due to the action of asymptomatic decompression gasformation and acute decompression illness of mild degree are presented. The authors propose the system of measures that would help to decrease the incidence of disease and to make the treatment of decompression illness more effective.

Acute Disease↗

Physiological control of diving behaviour in the Weddell seal Leptonychotes weddelli: a model based on cardiorespiratory control theory.

Despite being obligate air breathers, many species of marine mammal are capable of spending most of their lives submerged in water. How they do this has been a subject of intense interest to physiologists for over a century, yet we still do not have a detailed understanding of the physiological mechanisms underlying this behaviour. What are the proximate mechanisms that trigger the 'decisions' to submerge and return to the surface? The present study proposes a model intended to address this question, based on fundamental concepts of cardiorespiratory control. Two basic hypotheses are examined by computer simulation, using a mathematical model of the mammalian cardiorespiratory control system with parameter values for an adult Weddell seal: (1) that the control of diving can be considered to be a respiratory control problem, and (2) that dives are initiated and maintained by disfacilitation of respiratory drive, not inhibition. Computer simulations confirmed the plausibility of these hypotheses. Simulated diving behaviour and physiological responses (ventilation, cardiac output, blood and tissue gas tensions) were consistent with published data from freely diving Weddell seals. Dives up to the estimated aerobic dive limit (ADL, 18-25 min) could be simulated without the need for active inhibition of breathing in this model. This theoretical analysis suggests that the most important physiological adjustments occur during the surface interval phase of the dive cycle and include hyperventilation accompanied by high cardiac output, appropriate regulation of cerebral blood flow and central chemoreceptor threshold shifts. During dives, cardiac output, distribution of peripheral blood flow, splenic contraction and peripheral chemoreflex drives were found to modulate physiological and behavioural responses, but were not essential for simulated dives to occur. The main conclusion from this study is that the central chemoreceptor may be an important mechanism involved in the regulation of diving behaviour, implying that CO2, not O2, is the key regulatory variable in this model. This model includes and extends the ADL concept and suggests an explicit mechanism by which the respiratory control system may play a central role in the regulation of diving behaviour. It is likely that respiratory mechanisms are an important component of a hierarchical behavioural control system and further studies are required to test the qualitative and quantitative validity of the model.

Animals↗

Diurnal and seasonal variations in the duration and depth of the longest dives in southern elephant seals (Mirounga leonina): possible physiological and behavioural constraints.

This study seeks to understand how the physiological constraints of diving may change on a daily and seasonal basis. Dive data were obtained from southern elephant seals (Mirounga leonina) from South Georgia using satellite relay data loggers. We analysed the longest (95th percentile) dive durations as proxies for physiological dive limits. A strong, significant relationship existed between the duration of these dives and the time of day and week of year in which they were performed. The depth of the deepest dives also showed a significant, but far less consistent, relationship with local time of day and season. Changes in the duration of the longest dives occurred irrespective of their depth. Dives were longest in the morning (04:00-12:00 h) and shortest in the evening (16:00-00:00 h). The size of the fluctuation varied among animals from 4.0 to 20.0 min. The daily pattern in dive depth was phase-shifted in relation to the diurnal rhythm in dive duration. Dives were deeper at midday and shallower around midnight. Greater daily changes in duration occurred in seals feeding in the open ocean than in those foraging on the continental shelf. The seasonal peak in the duration of the longest dives coincided with austral midwinter. The size of the increase in dive duration from autumn/spring to winter ranged from 11.5 to 30.0 min. Changes in depth of the longest dives were not consistently associated with particular times of year. The substantial diurnal and seasonal fluctuations in maximum dive duration may be a result of changes in the physiological capacity to remain submerged, in addition to temporal changes in the ecological constraints on dive behaviour. We speculate about the role of melatonin as a hormonal mediator of diving capability.

Animals↗

The influence of body size on the diving behaviour and physiology of the bimodally respiring turtle, Elseya albagula.

In aquatic vertebrates that acquire oxygen aerially dive duration scales positively with body mass, i.e. larger animals can dive for longer periods, however in bimodally respiring animals the relationship between dive duration and body mass is unclear. In this study we investigated the relationships between body size, aquatic respiration, and dive duration in the bimodally respiring turtle, Elseya albagula. Under normoxic conditions, dive duration was found to be independent of body mass. The dive durations of smaller turtles were equivalent to that of larger individuals despite their relatively smaller oxygen stores and higher mass specific metabolic rates. Smaller turtles were able to increase their dive duration through the use of aquatic respiration. Smaller turtles had a relatively higher cloacal bursae surface area than larger turtles, which allowed them to extract a relatively larger amount of oxygen from the water. By removing the ability to respire aquatically (hypoxic conditions), the dive duration of the smaller turtles significantly decreased restoring the normal positive relationship between body size and dive duration that is seen in other air-breathing vertebrates.

Animals↗