Dive response of children in relation to cold-water near-drowning

The strength of the dive response, as judged by the combination of breath-hold duration (BHD) and bradycardia, was compared in 87 children (4-13 yr old) and 68 adults (20-68 yr old) during simulated dives in 29 degrees C water. Mean BHD in children (16.1 s) was only 37.4% (P less than 0.001) of adult BHD (43.0 s). Within children, BHD was significantly (P less than 0.001) dependent on age (A in yr) according to the regression BHD = -1.46 + 2.27A. No age dependency of BHD occurred in adults. Due to the low BHD of children, only 14/87 (16.1%) were able to breath hold for the 25 s necessary to develop full diving bradycardia. For these 14 children, their bradycardia (36.1% reduction) was insignificantly different (P greater than 0.50) from that of adults (36.4%). These experimental findings demonstrate that the dive response of children is extremely weak, due mainly to their very low BHD. Since lower water temperature would probably accentuate the shortness of BHD (according to previous findings for adults), it is concluded that the dive response is unlikely to make a significant contribution to the prolonged resuscitatibility of children who are victims of cold-water near-drowning.     

Thermal balance and survival time prediction of man in cold water.

Metabolic rates and rectal temperatures were continuously monitored for humans immersed in cold ocean water (4.6--18.2 degrees C) under stimulated accident conditions. The subjects wore only light clothing and a kapok lifejacket while either holding-still or swimming. While holding-still, metabolic heat production (Hm,kcal-min--1) was inversely related to water temperature (Tw, degrees C) according to the equation Hm equals 4.19 minus-0.117 Tw. This temperature response pattern is shown to be similar to that for exposure to air of the same temperature when air velocity is just over 5 m.p.h. (2.24 m/s). The thermogenic response was one-third efficient in balancing the calculated heat loss in cold water, resulting in hypothermia at a rectal temperature cooling rate (C, degrees C-min--1) dependent on water temperature (Tw, degrees C) according to the relation C equal 0.0785 - 0.0034Tw. Although swimming increased heat production to 2.5 times that of holding-still at 10.5 degrees C water temperature, cooling rate was 35% greater while swimming. A prediction equation for survival time (ts, min) of persons accidentally immersed in cold water (Tw, degrees C) has the form ts equal 15 + 7.2/(0.0785-0.0034Tw), based on the findings of this study, and it is compared to pre-existing models.     

Diving heart rate development in postnatal harbour seals, Phoca vitulina

Harbour seals, Phoca vitulina, dive from birth, providing a means of mapping the development of the diving response, and so our objective was to investigate the postpartum development of diving bradycardia. The study was conducted May-July 2000 and 2001 in the St. Lawrence River Estuary (48 degrees 41'N, 68 degrees 01'W). Both depth and heart rate (HR) were remotely recorded during 86,931 dives (ages 2-42 d, n = 15) and only depth for an additional 20,300 dives (combined data covered newborn to 60 d, n = 20). The mean dive depth and mean dive durations were conservative during nursing (2.1 +/- 0.1 m and 0.57 +/- 0.01 min, range = 0-30.9 m and 0-5.9 min, respectively). The HR of neonatal pups during submersion was bimodal, but as days passed, the milder of the two diving HRs disappeared from their diving HR record. By 15 d of age, most of the dive time was spent at the lower diving bradycardia rate. Additionally, this study shows that pups are born with the ability to maintain the lower, more fully developed dive bradycardia during focused diving but do not do so during shorter routine dives.     

Adaptive changes in the cardiac automatism of the sea otter Enhydra lutris

Electrographic investigations in sea otters reveal acute decrease in the heart rate (to 1/2--1/8 of the normal rate) during diving. The longer the dive, the more intense the bradycardia. Each partial exhalation under water causes further decrease in the heart rate. The duration of cardiac cycles (R--R) increases mainly at the expense of the prolongation of diastole (T--P). Variations in the heart rate during submersion result from the increased vagal tone. Atropine injections abolish diving bradycardia. On surfacing, the animals exhibit a pronounced recovery tachycardia.     

Body cooling and the diving capabilities of muskrats (Ondatra zibethicus): a test of the adaptive hypothermia hypothesis

We tested the hypothesis that immersion hypothermia enhances the diving capabilities of adult and juvenile muskrats by reducing rates of oxygen consumption (V O2). Declines in abdominal body temperature (T(b)) comparable to those observed in nature (0.5-3.5 degrees C) were induced by pre-chilling animals in 6 degrees C water. Pre-chilling did not reduce diving V O2 of any animal tested in 10 degrees C or 30 degrees C water, irrespective of the nature of the dive. Most behavioural indices of dive performance, including average and cumulative dive times, were unaffected by T(b) reduction in adults, but depressed in hypothermic juveniles (200-400 g). Hypothermia reduced diving heart rate only on short (<25s) dives (16% reduction, P=0.01), but did not affect the temporal onset of diving bradycardia. Post-immersion V O2 was higher for pre-chilled than for normothermic muskrats, but the difference became insignificant on longer (>90 s) dives. Our findings suggest that the mild hypothermia experienced by muskrats in nature has minimal effect on diving and post-immersion metabolic costs, and thus has little impact on the dive performance of this northern semi-aquatic mammal.     

Thermal plasticity of diving behavior, aquatic respiration, and locomotor performance in the Mary River turtle Elusor macrurus

Locomotion is a common measure of performance used in studies of thermal acclimation because of its correlation with predator escape and prey capture. However, for sedentary animals such as freshwater turtles, we propose that diving behavior may be a more ecologically relevant measure of performance. Increasing dive duration in hatchling turtles reduces predator exposure and therefore functions as an ecological benefit. Diving behavior is thermally dependent, and in some species of freshwater turtles, it is also reliant on aquatic respiration. This study examined the influence of thermal acclimation on diving behavior, aquatic respiration, and locomotor performance in the endangered, bimodally respiring Mary River turtle Elusor macrurus. Diving behavior was found to partially acclimate at 17 degrees C, with turtles acclimated to a cold temperature (17 degrees C) having a significantly longer dive duration than hatchlings acclimated to a warm temperature (28 degrees C). This increase in dive duration at 17 degrees C was not a result of physiological alterations in metabolic rate but was due instead to an increase in aquatic oxygen consumption. Increasing aquatic oxygen consumption permitted cold-acclimated hatchlings to remain submerged for significantly longer periods, with one turtle undertaking a dive of over 2.5 d. When burst-swimming speed was used as the measure of performance, thermal acclimation was not detected. Overall, E. macrurus demonstrated a partial ability to acclimate to changes in environmental temperature.     

The effect of clothing on "diving bradycardia" in man during submersion in cold water

Eighteen healthy male volunteers undertook three seated submersions into stirred water at 5 degrees C. Whilst submerged, the subjects attempted to hold their breath for 20 s. They wore a different clothing assembly for each submersion, viz: a cotton overall assembly, a "wet suit" assembly and a "dry suit" assembly. During the experiments the breath-hold time, heart rate, skin and rectal temperatures of the subjects were recorded. The results showed that significantly (P less than 0.05) more subjects developed a diving bradycardia--defined as five or more consecutive R-R intervals of over 1.2 s--when wearing the dry suit. It is concluded that increasing the cold stress experienced by individuals during cold-water submersion decreases the incidence of diving bradycardia but not the magnitude of the bradycardia when it occurs.     

Recording the free-living behaviour of small-bodied, shallow-diving animals with data loggers

1. Time-depth data recorders (TDRs) have been widely used to explore the behaviour of relatively large, deep divers. However, little is known about the dive behaviour of small, shallow divers such as semi-aquatic mammals. 2. We used high-resolution TDRs to record the diving behaviour of American mink Mustela vison (weight of individuals 580-1275 g) in rivers in Oxfordshire (UK) between December 2005 and March 2006. 3. Dives to > 0.2 m were measured in all individuals (n = 6). Modal dive depth and duration were 0.3 m and 10 s, respectively, although dives up to 3 m and 60 s in duration were recorded. Dive duration increased with dive depth. 4. Temperature data recorded by TDRs covaried with diving behaviour: they were relatively cold (modal temperature 4-6 degrees C across individuals) when mink were diving and relatively warm (modal temperature 24-36 degrees C across individuals) when mink were not diving. 5. Individuals differed hugely in their use of rivers, reflecting foraging plasticity across both terrestrial and aquatic environments. For some individuals there was < 1 dive per day while for others there was > 100 dives per day. 6. We have shown it is now possible to record the diving behaviour of small free-living animals that only dive a few tens of centimetres, opening up the way for a new range of TDR studies on shallow diving species.     

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.     

Diving through the thermal window: implications for a warming world

Population decline and a shift in the geographical distribution of some ectothermic animals have been attributed to climatic warming. Here, we show that rises in water temperature of a few degrees, while within the thermal window for locomotor performance, may be detrimental to diving behaviour in air-breathing ectotherms (turtles, crocodilians, marine iguanas, amphibians, snakes and lizards). Submergence times and internal and external body temperature were remotely recorded from freshwater crocodiles (Crocodylus johnstoni) while they free-ranged throughout their natural habitat in summer and winter. During summer, the crocodiles'' mean body temperature was 5.2 ± 0.1°C higher than in winter and the largest proportion of total dive time was composed of dive durations approximately 15 min less than in winter. Diving beyond 40 min during summer required the crocodiles to exponentially increase the time they spent on the surface after the dive, presumably to clear anaerobic debt. The relationship was not as significant in winter, even though a greater proportion of dives were of a longer duration, suggesting that diving lactate threshold (DLT) was reduced in summer compared with winter. Additional evidence for a reduced DLT in summer was derived from the stronger influence body mass exerted upon dive duration, compared to winter. The results demonstrate that the higher summer body temperature increased oxygen demand during the dive, implying that thermal acclimatization of the diving metabolic rate was inadequate. If the study findings are common among air-breathing diving ectotherms, then long-term warming of the aquatic environment may be detrimental to behavioural function and survivorship.     

Water temperature sampling by foraging Brünnich's Guillemots with bird-borne data loggers

We describe the features of waters where seabirds were feeding by sampling vertical water temperature profiles with data loggers mounted on five Brünnich's Guillemots in Svalbard, Norway. The guillemots foraged in a cold water (−0.5–0.5°C SST (sea surface temperature)) by making 1.8 dive bouts in short trips (32–257 min duration) as well as in moderate (0.5–2.0°C SST) and warm waters (2.5–4.0°C SST) by making 6.0 dive bouts during long trips (411–688 min duration). Judging from outbound flying time (15.7–24.4 min), time between dive bouts (23.9–43.3 min) and water types, the birds probably fed in fjord or coastal waters during short trips and in both coastal and offshore waters during long trips. Water temperature and diving behaviour can be simultaneously recorded by small data loggers, which therefore will provide useful information on marine features and foraging activity of top predators.     

Estimating the rate of oxygen consumption during submersion from the heart rate of diving animals

How animals manage their oxygen stores during diving and other breath-hold activities has been a topic of debate among physiologists for decades. Specifically, while the behavior of free-ranging diving animals suggests that metabolism during submersion must be primarily aerobic in nature, no studies have been able to determine their rate of oxygen consumption during submersion (Vo(2)d) and hence prove that this is the case. In the present study, we combine two previously used techniques and develop a new model to estimate Vo(2)d accurately and plausibly in a free-ranging animal and apply it to data for macaroni penguins (Eudyptes chrysolophus) as an example. For macaroni penguins at least, Vo(2)d can be predicted by measuring heart rate during the dive cycle and the subsequent surface interval duration. Including maximum depth of the dive improves the accuracy of these predictions. This suggests that energetically demanding locomotion events within the dive combine with the differing buoyancy and locomotion costs associated with traveling to depth to influence its cost in terms of oxygen use. This will in turn effect the duration of the dive and the duration of the subsequent recovery period. In the present study, Vo(2)d ranged from 4 to 28 ml.min(-1).kg(-1), indicating that, at least as far as aerobic metabolism was concerned, macaroni penguins were often hypometabolic, with rates of oxygen consumption usually below that for this species resting in water (25.6 ml.min(-1).kg(-1)) and occasionally lower than that while resting in air (10.3 ml.min(-1).kg(-1)).     

Body temperature and insulation in diving Great Cormorants and European Shags

1. Cormorants are typically considered as wettable diving birds with high thermoregulatory costs and are presumed to exert substantial predatory pressure on fish stocks.
2. The stomach temperatures of seven Great Cormorants and three European Shags were recorded during a total of 108 foraging trips undertaken near the Chausey Islands breeding colony (France).
3. Both species kept a constant body temperature during the dive series which lasted up to 158 min and were conducted in 12°C water. Consequently, assuming that heat loss to the water is equal to heat production in diving Great Cormorants, the minimal insulating plumage air volume was calculated to be 0·371 × 10^–3 m³ (corresponding to a 1·62-mm air layer) in males and 0·347 × 10^–3 m³ (corresponding to a 1·90-mm air layer) in females.
4. Furthermore, it is shown that plumage air volume and dive depth are the major factors influencing heat flux to the water and that the energetics of diving Great Cormorants may also vary substantially according to fat layer thickness, water temperature and body temperature. Swim speed plays only a minor role.
5. Considering these results, it is postulated that Great Cormorants may have optimized plumage air volume so as to minimize both mechanical costs (upthrust) and thermoregulatory costs of swimming in cold, shallow water.
6. Finally, body temperature patterns recorded in different cormorant species while diving are compared.     

Effects of diving and swimming behavior on body temperatures of pacific leatherback turtles in tropical seas

Mathematical models and recordings of cloacal temperature suggest that leatherback turtles (Dermochelys coriacea) maintain core body temperature higher than ambient water temperature (T(W)) while freely swimming at sea. We investigated the thermoregulatory capabilities of free-ranging leatherbacks and, specifically, the effect that changes in diving patterns and ambient temperatures have on leatherback body temperatures (T(B)). Data loggers were used to record subcarapace and gastrointestinal tract temperatures (T(SC) and T(GT), respectively), T(W), swim speed, dive depth, and dive times of female leatherback turtles during internesting intervals off the coast of Guanacaste, Costa Rica. Mean T(SC) (28.7 degrees -29.0 degrees C) was significantly higher than mean T(W) (25.0 degrees -27.5 degrees C). There was a significant positive relationship between T(SC) and T(W) and a significant negative correlation between T(SC) and dive depth and T(GT) and dive depth. Rapid fluctuations in T(GT) occurred during the first several days of the internesting interval, which suggests that turtles were ingesting prey or water during this time. Turtles spent 79%-91% of the time at sea swimming at speeds greater than 0.2 m s(-1), and the average swim speed was 0.7 +/- 0.2 m s(-1). Results from this study show that alterations in diving behavior and T(W) affect T(B) of leatherback turtles in the tropics. Body temperatures of free-ranging leatherback turtles correspond well with values for T(B) predicted by mathematical models for tropical conditions.     

In vivo thermal conductivity of the human forearm tissues

The effective thermal conductivities of the skin + subcutaneous (keff skin + fat) and muscle (keff muscle) tissues of the human forearm at thermal steady state during immersion in water at temperatures (Tw) ranging from 15 to 36 degrees C were determined. Tissue temperature (Tt) was continuously monitored by a calibrated multicouple probe during a 3-h immersion of the resting forearm. Tt was measured every 5 mm from the longitudinal axis of the forearm (determined from computed-tomography scanning) to the skin surface. Skin temperature (Tsk), heat loss (Hsk), and blood flow (Q) of the forearm, as well as rectal temperature (Tre) and arterial blood temperature at the brachial artery (Tbla), were measured during the experiments. When the keff values were calculated from the finite-element (FE) solution of the bioheat equation, keff skin + fat ranged from 0.28 +/- 0.03 to 0.73 +/- 0.14 W.degrees C-1.m-1 and keff muscle varied between 0.56 +/- 0.05 and 1.91 +/- 0.19 W.degrees C-1.m-1 from 15 to 36 degrees C. The values of keff skin + fat and keff muscle, calculated from the FE solution for Tw less than or equal to 30 degrees C, were not different from the average in vitro values obtained from the literature. The keff values of the forearm tissues were linearly related (r = 0.80, P less than 0.001) to Q for Tw greater than or equal to 30 degrees C. It was found that the muscle tissue could account for 92 +/- 1% of the total forearm insulation during immersion in water between 15 and 36 degrees C.     

The influence of temperature on diving behaviour in the alpine newt, Triturus alpestris

An aquatic lifestyle poses serious restriction to air-breathing animals in terms of time and energy spent during a dive cycle. The diving frequency increases with water temperature, therefore an ectotherm's time budget greatly depends on the thermal characteristics of the aquatic environment. Available data suggests that time costs caused by temperature-dependent dive frequency can be partially compensated for by adjusting the swimming speed and diving angle during dive cycle. We tested this prediction by examining the influence of temperature on the diving behaviour of the alpine newt, Triturus alpestris. The ascending speed and angle showed disparate patterns of temperature dependency, with a minor influence on travel duration. Surprisingly, at higher temperatures, the diving newts saved most of their time by restricting swimming activity in the water column during their return to the bottom and not by adjusting their ascending duration. Hence, aquatic newts have the capacity to reduce temperature-dependent time costs of aerial breathing primarily by behavioural modifications during the descending phase of the dive cycle.     

Diving behaviour of a reptile (Crocodylus johnstoni) in the wild: interactions with heart rate and body temperature

The differences in physical properties of air and water pose unique behavioural and physiological demands on semiaquatic animals. The aim of this study was to describe the diving behaviour of the freshwater crocodile Crocodylus johnstoni in the wild and to assess the relationships between diving, body temperature, and heart rate. Time-depth recorders, temperature-sensitive radio transmitters, and heart rate transmitters were deployed on each of six C. johnstoni (4.0-26.5 kg), and data were obtained from five animals. Crocodiles showed the greatest diving activity in the morning (0600-1200 hours) and were least active at night, remaining at the water surface. Surprisingly, activity pattern was asynchronous with thermoregulation, and activity was correlated to light rather than to body temperature. Nonetheless, crocodiles thermoregulated and showed a typical heart rate hysteresis pattern (heart rate during heating greater than heart rate during cooling) in response to heating and cooling. Additionally, dive length decreased with increasing body temperature. Maximum diving length was 119.6 min, but the greatest proportion of diving time was spent on relatively short (<45 min) and shallow (<0.4 m) dives. A bradycardia was observed during diving, although heart rate during submergence was only 12% lower than when animals were at the surface.     

Forearm temperature profile during the transient phase of thermal stress.

The transient temperature response of the resting human forearm immersed in water at temperatures (Tw) ranging from 15 to 36 degrees C was investigated. Tissue temperature (Tt) was continuously monitored by a calibrated multicouple probe during the 3-h immersions. Tt was measured every 5 mm, from the longitudinal axis of the forearm to the skin surface. Skin temperature, rectal temperature, and blood flow (Q) were also measured during the immersions. The maximum rate of change of the forearm mean tissue temperature (Tt, max) occurred during the first 5 min of the immersion. Tt, max was linearly dependent on Tw (P less than 0.001), with mean values (SEM) ranging from -0.8 (0.1) degrees C.min-1 at 15 degrees C to 0.2 (0.1) degrees C.min-1 at 36 degrees C. The maximum rate of change of compartment mean temperature was dependent (P less than 0.001) on the radial distance from the longitudinal axis of the forearm. The half-time for thermal steady state of the forearm mean tissue temperature was linearly dependent on Tw between 30 and 36 degrees C (P less than 0.01), with mean values (SEM) ranging from 15.6 (0.6) min at 30 degrees C to 9.7 (1.2) min at 36 degrees C and not different between 15 and 30 degrees C, averaging 16.2 (0.6) min. There was a significant linear relationship between the half-time for thermal steady-state of the compartment mean temperature and the radial distance from the longitudinal axis of the forearm for each value of Tw tested (P less than 0.001). The data of the present study suggest that the forearm Q is an important determinant of the transient thermal response of the forearm tissue during thermal stress.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

Diving in shallow water: the foraging ecology of darters (Aves: Anhingidae)

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.