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It is fundamentally important for many animal ecologists to quantify the costs of animal activities, although it is not straightforward to do so. The recording of triaxial acceleration by animal-attached devices has been proposed as a way forward for this, with the specific suggestion that dynamic body acceleration (DBA) be used as a proxy for movement-based power. Dynamic body acceleration has now been validated frequently, both in the laboratory and in the field, although the literature still shows that some aspects of DBA theory and practice are misunderstood. Here, we examine the theory behind DBA and employ modelling approaches to assess factors that affect the link between DBA and energy expenditure, from the deployment of the tag, through to the calibration of DBA with energy use in laboratory and field settings. Using data from a range of species and movement modes, we illustrate that vectorial and additive DBA metrics are proportional to each other. Either can be used as a proxy for energy and summed to estimate total energy expended over a given period, or divided by time to give a proxy for movement-related metabolic power. Nonetheless, we highlight how the ability of DBA to predict metabolic rate declines as the contribution of non-movement-related factors, such as heat production, increases. Overall, DBA seems to be a substantive proxy for movement-based power but consideration of other movement-related metrics, such as the static body acceleration and the rate of change of body pitch and roll, may enable researchers to refine movement-based metabolic costs, particularly in animals where movement is not characterized by marked changes in body acceleration. Fil: Wilson, Rory P.. Swansea University; Reino Unido Fil: Börger, Luca. Swansea University; Reino Unido Fil: Holton, Mark. Swansea University; Reino Unido Fil: Scantlebury, D. Michael. The Queens University of Belfast; Irlanda Fil: Gómez Laich, Agustina Marta. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina Fil: Quintana, Flavio Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina Fil: Rosell, Frank. University of South‐Eastern Norway; Noruega Fil: Graf, Patricia M.. Universitat Fur Bodenkultur Wien; Austria. University of Ljubljana; Eslovenia Fil: Williams, Hannah. Swansea University; Reino Unido Fil: Gunner, Richard. Swansea University; Reino Unido Fil: Hopkins, Lloyd. Swansea University; Reino Unido Fil: Marks, Nikki. The Queens University of Belfast; Irlanda Fil: Geraldi, Nathan R.. King Abdullah University of Science and Technology; Arabia Saudita Fil: Duarte, Carlos M.. King Abdullah University of Science and Technology; Arabia Saudita Fil: Scott, Rebecca. Geomar-Helmholtz Centre for Ocean Research Kiel; Alemania Fil: Strano, Michael S.. Massachusetts Institute of Technology; Estados Unidos Fil: Robotka, Hermina. Max Planck Institute für Ornithologie; Alemania Fil: Eizaguirre, Christophe. , Queen Mary University of London; Reino Unido Fil: Fahlman, Andreas. undación Oceanogràfic de la Comunidad Valenciana; España Fil: Shepard, Emily L. C.. Swansea University; Reino Unido. Max Planck Institute für Ornithologie; Alemania

In the current study, we used breath-by-breath respirometry to evaluate respiratory physiology under voluntary control in a male beluga calf [Delphinapterus leucas, body mass range (M b): 151-175 kg], an adult female (estimated M b = 500-550 kg) and a juvenile male (M b = 279 kg) false killer whale (Pseudorca crassidens) housed in managed care. Our results suggest that the measured breathing frequency (f R) is lower, while tidal volume (V T) is significantly greater as compared with allometric predictions from terrestrial mammals. Including previously published data from adult bottlenose dolphin (Tursiops truncatus) beluga, harbor porpoise (Phocoena phocoena), killer whale (Orcinus orca), pilot whale (Globicephala scammoni), and gray whale (Eschrichtius robustus) show that the allometric mass-exponents for V T and f R are similar to that for terrestrial mammals (V T: 1.00, f R: -0.20). In addition, our results suggest an allometric relationship for respiratory flow ( V . ), with a mass-exponent between 0.63 and 0.70, and where the expiratory V . was an average 30% higher as compared with inspiratory V . . These data provide enhanced understanding of the respiratory physiology of cetaceans and are useful to provide proxies of lung function to better understand lung health or physiological limitations.

Theoretical models are used to predict how breath-hold diving vertebrates manage O2, CO2, and N2 while underwater. One recent gas dynamics model used available lung and tracheal compliance data from various species. As variation in respiratory compliance significantly affects alveolar compression and pulmonary shunt, the current study objective was to evaluate changes in model output when using species-specific parameters from California sea lions (Zalophus californianus). We explored the effects of lung and dead space compliance on the uptake of N2, O2, and CO2 in various tissues during a series of hypothetical dives. The updated parameters allowed for increased compliance of the lungs and an increased stiffness in the trachea. When comparing updated model output with a model using previous compliance values, there was a large decrease in N2 uptake but little change in O2 and CO2 levels. Therefore, previous models may overestimate N2 tensions and the risk of gas-related disease, such as decompression sickness (DCS), in marine mammals.

Assessment of the compressibility of marine mammal airways at depth is crucial to understanding vital physiologic processes such as gas exchange during diving. Very few studies have directly assessed changes in cetacean and pinniped trachea-bronchial shape, and none have quantified changes in volume with increasing pressure. A freshly deceased harbor seal, grey seal, harp seal, harbor porpoise, and common dolphin were imaged post mortem via CT in a radiolucent hyperbaric chamber as previously described in Moore et al (2011). Volume reconstructions were performed of segments of the trachea and bronchi of the pinnipeds and bronchi of the cetaceans for each pressure treatment. All specimens examined demonstrated significant decreases in volume with increasing pressure, with the harbor seal and common dolphin nearing complete collapse at the highest pressures. The common dolphin bronchi demonstrated distinctly different compression dynamics between 50% and 100% lung inflation treatments, indicating the importance of air in maintaining patent airways, and collapse occurred caudally to cranially in the 50% treatment. Dynamics of the harbor seal and grey seal airways indicated that the trachea was less compliant than the bronchi. These findings indicate potential species-specific variability in airway compliance, and cessation of gas exchange may occur at greater depths than those predicted in models assuming rigid airways. This may potentially increase the likelihood of decompression sickness in these animals during diving.

The outstanding properties of graphene have initiated myriads of research and development; yet, its economic impact is hampered by the difficulties encountered in production and practical application. Recently discovered laser-induced graphene is generated by a simple printing process on flexible and lightweight polyimide films. Exploiting the electrical features and mechanical pliability of LIG on polyimide, we developed wearable resistive bending sensors that pave the way for many cost-effective measurement systems. The versatile sensors we describe can be utilized in a wide range of configurations, including measurement of force, deflection, and curvature. The deflection induced by different forces and speeds is effectively sensed through a resistance measurement, exploiting the piezoresistance of the printed graphene electrodes. The LIG sensors possess an outstanding range for strain measurements reaching >10% A double-sided electrode concept was developed by printing the same electrodes on both sides of the film and employing difference measurements. This provided a large bidirectional bending response combined with temperature compensation. Versatility in geometry and a simple fabrication process enable the detection of a wide range of flow speeds, forces, and deflections. The sensor response can be easily tuned by geometrical parameters of the bending sensors and the LIG electrodes. As a wearable device, LIG bending sensors were used for tracking body movements. For underwater operation, PDMS-coated LIG bending sensors were integrated with ultra-low power aquatic tags and utilized in underwater animal speed monitoring applications, and a recording of the surface current velocity on a coral reef in the Red Sea. Printed laser induced graphene (LIG) enables versatile multiple-parameter piezoresistive sensing in actual fields. An international team led by Prof Jurgen Kosel from King Abdullah University of Science and Technology, Saudi Arabia develops bending piezoresistive sensors based on laser induced graphene (LIG) on flexible polyimide substrate to accurately detect curvature, force, deflection and flow. When combined with double-sided device design to complete compensate the temperature effect and commercial ultra-low power aquatic tags, the LIG sensors can be used for various marine life measurements and monitoring, such as marine animals’ speed monitoring under realistic and harsh conditions. These LIG sensors presented here fulfill several key requirements such as low cost, light weight, flexibility and minimal intrusiveness and are thus promising for future applications in wearable electronics.

The dive response is well documented for marine mammals, and includes a significant reduction in heart rate (fH) during submersion as compared while breathing at the surface. In the current study we assessed the influence of the Respiratory Sinus Arrhythmia (RSA) while estimating the resting fH while breathing. Using transthoracic echocardiography we measured fH, and stroke volume (SV) during voluntary surface apneas at rest up to 255 s, and during recovery from apnea in 11 adult bottlenose dolphins (Tursiops truncatus, 9 males and 2 females, body mass range: 140–235 kg). The dolphins exhibited a significant post-respiratory tachycardia and increased SV. Therefore, only data after this RSA had stabilized were used for analysis and comparison. The average (±s.d.) fH, SV, and cardiac output (CO) after spontaneous breaths while resting at the surface were 44 ± 6 beats min−1, 179 ± 31 ml, and 7909 ± 1814 l min−1, respectively. During the apnea the fH, SV, and CO decreased proportionally with the breath-hold duration, and after 255 s they, respectively, had decreased by an average of 18%, 1–21%, and 12–37%. During recovery, the fH, SV, and CO rapidly increased by as much as 117%, 34%, and 190%, respectively. Next, fH, SV and CO rapidly decreased to resting values between 90–110 s following the surface apnea. These data highlight the necessity to define how the resting fH is estimated at the surface, and separating it from the RSA associated with each breath to evaluate the significance of cardiorespiratory matching during diving.

Mass stranding events (MSEs) of beaked whales (BWs) were extremely rare prior to the 1960s but increased markedly after the development of naval mid-frequency active sonar (MFAS). The temporal and spatial associations between atypical BW MSEs and naval exercises were first observed in the Canary Islands, Spain, in the mid-1980s. Further research on BWs stranded in association with naval exercises demonstrated pathological findings consistent with decompression sickness (DCS). A 2004 ban on MFASs around the Canary Islands successfully prevented additional BW MSEs in the region, but atypical MSEs have continued in other places of the world, especially in the Mediterranean Sea, with examined individuals showing DCS. A workshop held in Fuerteventura, Canary Islands, in September 2017 reviewed current knowledge on BW atypical MSEs associated with MFAS. Our review suggests that the effects of MFAS on BWs vary among individuals or populations, and predisposing factors may contribute to individual outcomes. Spatial management specific to BW habitat, such as the MFAS ban in the Canary Islands, has proven to be an effective mitigation tool and mitigation measures should be established in other areas taking into consideration known population-level information.

We measured respiratory flow ( V ), breathing frequency ( f R ), tidal volume ( V T ), breath duration and end-expired O 2 content in bottlenose dolphins ( Tursiops truncatus ) before and after static surface breath-holds ranging from 34 to 292 s. There was considerable variation in the end-expired O 2 , V T and f R following a breath-hold. The analysis suggests that the dolphins attempt to minimize recovery following a dive by altering V T and f R to rapidly replenish the O 2 stores. For the first breath following a surface breath-hold, the end-expired O 2 decreased with dive duration, while V T and f R increased. Throughout the recovery period, end-expired O 2 increased while the respiratory effort ( V T , f R ) decreased. We propose that the dolphins alter respiratory effort following a breath-hold according to the reduction in end-expired O 2 levels, allowing almost complete recovery after 1.2 min.

Instrumenting animals with tags contributes additional resistive forces (weight, buoyancy, lift, and drag) that may result in increased energetic costs; however, additional metabolic expense can be moderated by adjusting behavior to maintain power output. We sought to increase hydrodynamic drag for near-surface swimming bottlenose dolphins, to investigate the metabolic effect of instrumentation. In this experiment, we investigate whether (1) metabolic rate increases systematically with hydrodynamic drag loading from tags of different sizes or (2) whether tagged individuals modulate speed, swimming distance, and/or fluking motions under increased drag loading. We detected no significant difference in oxygen consumption rates when four male dolphins performed a repeated swimming task, but measured swimming speeds that were 34% ( > 1 m s-1) slower in the highest drag condition. To further investigate this observed response, we incrementally decreased and then increased drag in six loading conditions. When drag was reduced, dolphins increased swimming speed (+1.4 m s-1; +45%) and fluking frequency (+0.28 Hz; +16%). As drag was increased, swimming speed (-0.96 m s-1; -23%) and fluking frequency (-14 Hz; 7%) decreased again. Results from computational fluid dynamics simulations indicate that the experimentally observed changes in swimming speed would have maintained the level of external drag forces experienced by the animals. Together, these results indicate that dolphins may adjust swimming speed to modulate the drag force opposing their motion during swimming, adapting their behavior to maintain a level of energy economy during locomotion. Summary Statement: Biologging and tracking tags add drag to study subjects. When wearing tags of different sizes, dolphins changed their swimming paths, speed, and movements to modulate power output and energy consumption.

The accurate estimation of field metabolic rates (FMR) in wild animals is a key component of bioenergetic models, and is important for understanding the routine limitations for survival as well as individual responses to disturbances or environmental changes. Several methods have been used to estimate FMR, including accelerometer-derived activity budgets, isotope dilution techniques, and proxies from heart rate. Counting the number of breaths is another method used to assess FMR in cetaceans, which is attractive in its simplicity and the ability to measure respiration frequency from visual cues or data loggers. This method hinges on the assumption that over time a constant tidal volume (VT) and O2 exchange fraction (ΔO2) can be used to predict FMR. To test whether this method of estimating FMR is valid, we measured breath-by-breath tidal volumes and expired O2 levels of bottlenose dolphins, and computed the O2 consumption rate (V̇O2) before and after a pre-determined duration of exercise. The measured V̇O2 was compared with three methods to estimate FMR. Each method to estimate V̇O2 included variable VT and/or ΔO2. Two assumption-based methods overestimated V̇O2 by 216-501%. Once the temporal changes in cardio-respiratory physiology, such as variation in VT and ΔO2, were taken into account, pre-exercise resting V̇O2 was predicted to within 2%, and post-exercise V̇O2 was overestimated by 12%. Our data show that a better understanding of cardiorespiratory physiology significantly improves the ability to estimate metabolic rate from respiratory frequency, and further emphasizes the importance of eco-physiology for conservation management efforts., Summary: Accounting for changes in tidal volume and gas exchange improves the ability to estimate field metabolic rate from respiratory frequency in cetaceans.

Man-made environmental change may have significant impact on apex predators, like marine mammals. Thus, it is important to assess the physiological boundaries for survival in these species, and assess how climate change may affect foraging efficiency and the limits for survival. In the current study, we investigated whether the respiratory sinus arrhythmia (RSA) could estimate tidal volume (VT) in resting bottlenose dolphins (Tursiops truncatus). For this purpose, we measured respiratory flow and electrocardiogram (ECG) in five adult bottlenose dolphins at rest while breathing voluntarily. Initially, an exponential decay function, using three parameters (baseline heart rate, the change in heart rate following a breath, and an exponential decay constant) was used to describe the temporal change in instantaneous heart rate following a breath. The three descriptors, in addition to body mass, were used to develop a Generalized Additive Model (GAM) to predict the inspired tidal volume (VTinsp). The GAM allowed us to predict VTinsp with an average ( ± SD) overestimate of 3 ± 2%. A jackknife sensitivity analysis, where 4 of the five dolphins were used to fit the GAM and the 5th dolphin used to make predictions resulted in an average overestimate of 2 ± 10%. Future studies should be used to assess whether similar relationships exist in active animals, allowing VT to be studied in free-ranging animals provided that heart rate can be measured.

We measured respiratory flow (

To provide new insight into the pathophysiological mechanisms underlying gas emboli (GE) in bycaught loggerhead sea turtles (

In recent decades, biologists have sought to tag animals with various sensors to study aspects of their behavior otherwise inaccessible from controlled laboratory experiments. Despite this, chemical information, both environmental and physiological, remains challenging to collect despite its tremendous potential to elucidate a wide range of animal behaviors. In this work, we explore the design, feasibility, and data collection constraints of implantable, near-infrared fluorescent nanosensors based on DNA-wrapped single-wall carbon nanotubes (SWNT) embedded within a biocompatible poly(ethylene glycol) diacrylate (PEGDA) hydrogel. These sensors are enabled by Corona Phase Molecular Recognition (CoPhMoRe) to provide selective chemical detection for marine organism biologging. Riboflavin, a key nutrient in oxidative phosphorylation, is utilized as a model analyte in in vitro and ex vivo tissue measurements. Nine species of bony fish, sharks, eels, and turtles were utilized on site at Oceanogràfic in Valencia, Spain to investigate sensor design parameters, including implantation depth, sensor imaging and detection limits, fluence, and stability, as well as acute and long-term biocompatibility. Hydrogels were implanted subcutaneously and imaged using a customized, field-portable Raspberry Pi camera system. Hydrogels could be detected up to depths of 7 mm in the skin and muscle tissue of deceased teleost fish ( Sparus aurata and Stenotomus chrysops) and a deceased catshark ( Galeus melastomus). The effects of tissue heterogeneity on hydrogel delivery and fluorescence visibility were explored, with darker tissues masking hydrogel fluorescence. Hydrogels were implanted into a living eastern river cooter ( Pseudemys concinna), a European eel ( Anguilla anguilla), and a second species of catshark ( Scyliorhinus stellaris). The animals displayed no observable changes in movement and feeding patterns. Imaging by high-resolution ultrasound indicated no changes in tissue structure in the eel and catshark. In the turtle, some tissue reaction was detected upon dissection and histopathology. Analysis of movement patterns in sarasa comet goldfish ( Carassius auratus) indicated that the hydrogel implants did not affect swimming patterns. Taken together, these results indicate that this implantable form factor is a promising technique for biologging using aquatic vertebrates with further development. Future work will tune the sensor detection range to the physiological range of riboflavin, develop strategies to normalize sensor signal to account for the optical heterogeneity of animal tissues, and design a flexible, wearable device incorporating optoelectronic components that will enable sensor measurements in moving animals. This work advances the application of nanosensors to organisms beyond the commonly used rodent and zebrafish models and is an important step toward the physiological biologging of aquatic organisms.

We investigated lung function in bycaught sea turtles with and without gas emboli. Hyperbaric O2 treatment significantly improves lung function. Pulmonary function testing may be a useful diagnostic tool to assess fisheries induced trauma., Fisheries interactions are the most serious threats for sea turtle populations. Despite the existence of some rescue centres providing post-traumatic care and rehabilitation, adequate treatment is hampered by the lack of understanding of the problems incurred while turtles remain entrapped in fishing gears. Recently it was shown that bycaught loggerhead sea turtles (Caretta caretta) could experience formation of gas emboli (GE) and develop decompression sickness (DCS) after trawl and gillnet interaction. This condition could be reversed by hyperbaric O2 treatment (HBOT). The goal of this study was to assess how GE alters respiratory function in bycaught turtles before recompression therapy and measure the improvement after this treatment. Specifically, we assessed the effect of DCS on breath duration, expiratory and inspiratory flow and tidal volume (VT), and the effectiveness of HBOT to improve these parameters. HBOT significantly increased respiratory flows by 32–45% while VT increased by 33–35% immediately after HBOT. Repeated lung function testing indicated a temporal increase in both respiratory flow and VT for all bycaught turtles, but the changes were smaller than those seen immediately following HBOT. The current study suggests that respiratory function is significantly compromised in bycaught turtles with GE and that HBOT effectively restores lung function. Lung function testing may provide a novel means to help diagnose the presence of GE, be used to assess treatment efficacy, and contribute to sea turtle conservation efforts.

We measured respiratory flow rates, and expired O2in 32 (2–34 years, body mass [Mb] range: 73–291 kg) common bottlenose dolphins (Tursiops truncatus) during voluntary breaths on land or in water (between 2014 and 2017). The data were used to measure the resting O2consumption rate (V˙O2, range: 0.76–9.45 ml O2 min−1 kg−1) and tidal volume (VT, range: 2.2–10.4 l) during rest. For adult dolphins, the restingVT, but notV˙O2, correlated with body mass (Mb, range: 141–291 kg) with an allometric mass-exponent of 0.41. These data suggest that the mass-specificVTof larger dolphins decreases considerably more than that of terrestrial mammals (mass-exponent: 1.03). The average restingsV˙O2was similar to previously published metabolic measurements from the same species. Our data indicate that the resting metabolic rate for a 150 kg dolphin would be 3.9 ml O2 min−1 kg−1, and the metabolic rate for active animals, assuming a multiplier of 3–6, would range from 11.7 to 23.4 ml O2 min−1 kg−1.\absbreak Our measurements provide novel data for resting energy use and respiratory physiology in wild cetaceans, which may have significant value for conservation efforts and for understanding the bioenergetic requirements of this species.

Marine mammals are repeatedly exposed to elevated extra-thoracic pressure and alveolar collapse during diving and readily experience alveolar expansion upon inhalation – a unique capability as compared to terrestrial mammals. How marine mammal lungs overcome the challenges of frequent alveolar collapse and recruitment remains unknown. Recent studies indicate that pinniped lung surfactant has more anti-adhesive components compared to terrestrial mammals, which would aid in alveolar opening. However, pulmonary surfactant composition has not yet been investigated in odontocetes, whose physiology and diving behavior differ from pinnipeds. The aim of this study was to investigate the phosphatidylcholine (PC) composition of lung surfactants from various marine mammals and compare these to a terrestrial mammal. We found an increase in anti-adhesive PC species in harp seal (Pagophilus groenlandicus) and California sea lion (Zalophus californianus) compared to dog (Canus lupus familiaris), as well as an increase in the fluidizing PCs 16:0/14:0 and 16:0/16:1 in pinnipeds compared to odontocetes. The harbor porpoise (a representative of the odontocetes) did not have higher levels of fluidizing PCs compared to dog. Our preliminary results support previous findings that pinnipeds may have adapted unique surfactant compositions that allow them to dive at high pressures for extended periods without adverse effects. Future studies will need to investigate the differences in other surfactant components to fully assess the surfactant composition in odontocetes.

Incidental capture, or ‘bycatch’ in fishing gear is a major global threat to sea turtle populations. A recent study showed that underwater entrapment in fishing gear followed by rapid decompression may cause gas bubble formation within the blood stream (embolism) and tissues leading to organ injury, impairment, and even mortality in some bycaught individuals. We analyzed data from 128 capture events using logistic and ordinal regression to examine risk factors associated with gas embolism in sea turtles captured in trawls and gillnets. Likelihood of fatal decompression increases with increasing depth of gear deployment. A direct relationship was found between depth, risk and severity of embolism, which has not been previously demonstrated in any breath-hold diving species. For the trawl fishery in this study, an average trawl depth of 65 m was estimated to result in 50% mortality in by-caught turtles throughout the year. This finding is critical for a more accurate estimation of sea turtle mortality rates resulting from different fisheries and for devising efforts to avoid or minimize the harmful effects of capture.

1. A reference to the metabolic cost in Patagonia sea lions is missing in the first paragraph of the discussion. The following sentence should be corrected: The estimated resting metabolic rates were similar to those measured in Steller sea lions and California sea lions in water (Hurley and Costa, 2001; Fahlman et al., 2008, 2013) and Steller sea lions in air (Rosen and Trites, 2000).

Swimming at an optimal speed is critical for breath-hold divers seeking to maximize the time they can spend foraging underwater. Theoretical studies have predicted that the optimal swim speed for an animal while transiting to and from depth is independent of buoyancy, but is dependent on drag and metabolic rate. However, this prediction has never been experimentally tested. Our study assessed the effects of buoyancy and drag on the swim speed of three captive Steller sea lions (Eumetopias jubatus) that made 186 dives. Our study animals were trained to dive to feed at fixed depths (10–50 m) under artificially controlled buoyancy and drag conditions. Buoyancy and drag were manipulated using a pair of polyvinyl chloride (PVC) tubes attached to harnesses worn by the sea lions, and buoyancy conditions were designed to fall within the natural range of wild animals (∼12–26% subcutaneous fat). Drag conditions were changed with and without the PVC tubes, and swim speeds were recorded and compared during descent and ascent phases using an accelerometer attached to the harnesses. Generalized linear mixed-effect models with the animal as the random variable and five explanatory variables (body mass, buoyancy, dive depth, dive phase, and drag) showed that swim speed was best predicted by two variables, drag and dive phase (AIC = −139). Consistent with a previous theoretical prediction, the results of our study suggest that the optimal swim speed of Steller sea lions is a function of drag, and is independent of dive depth and buoyancy.

Three Steller sea lions Eumetopias jubatus, trained to dive voluntarily to depths ranging from 10 to 50 m, were used to determine whether the relationship between activity and metabolic rate during a diving interval (MRDI, dive + surface interval) was affected by fasting (9 d) during the breeding season (spring through summer). We subsequently used the relationship between activity and MRDI to partition the metabolic costs between underwater breath-holding activity and surface breathing activities. We estimated activity from overall dynamic body accel- eration (ODBA) measured using a 3-axis accelerometer, and measured MRDI using flow-through respirometry. The relationship between ODBA-based activity and MRDI was not affected by fast- ing period, suggesting that ODBA can be used to predict energy expenditure regardless of nutri- tional state in the spring and summer. However, the relationship between ODBA and dive meta- bolic rate differs from the relationship between ODBA and the surface metabolic rate before diving. Partitioning MRDI into the metabolic cost of remaining at the surface versus swimming underwater suggests that the metabolic cost of diving for Steller sea lions is approximately 29% lower than when breathing at the surface. ODBA appears to be a reasonable proxy to estimate metabolic rate in marine mammals, but more detailed behavioral data may be required to accu- rately apply the method in the field.

We measured esophageal pressures (n = 4), respiratory flow rates (n = 5), and expired O2 and CO2 (n = 4) in five adult Patagonia sea lions (Otaria flavescens, body mass range 94.3–286.0 kg) during voluntary breaths while laying down out of water. The data were used to estimate the dynamic specific lung compliance (sCL), the O2 consumption rate (V˙O2) and CO2 production rates (V˙CO2) during rest. Our results indicate that the resting tidal volume in Patagonia sea lions is approximately 47–73% of the estimated total lung capacity. The esophageal pressures indicated that expiration is passive during voluntary breaths. The average sCL of sea lions was 0.41 ± 0.11 cmH2O−1, which is similar to those measured in anesthetized sea lions and awake cetaceans, and significantly higher as compared to humans (0.08 cmH2O−1). The average estimated V˙O2 and V˙CO2 using breath-by-breath respirometry were 1.023 ± 0.327 L O2 min−1 (range: 0.695–1.514 L O2 min−1) and 0.777 ± 0.318 L CO2 min−1, (range: 0.510–1.235 L CO2 min−1), respectively, which is similar to previously published metabolic measurements from California and Steller sea lions using conventional flow-through respirometry. Our data provide end-tidal gas composition and offer novel data for respiratory physiology in pinnipeds, which may be important for clinical medicine and conservation efforts.

A probabilistic model was used to predict decompression sickness (DCS) outcome in pig (70 and 20 kg), hamster (100 g), rat (220 g) and mouse (20 g) following air saturation dives. The data set included 179 pig, 200 hamster, 360 rat, and 224 mouse exposures to saturation pressures ranging from 1.9–15.2 ATA and with varying decompression rates (0.9–156 ATA • min−1). Single exponential kinetics described the tissue partial pressures (Ptiss) of N2: Ptiss = ∫(Pamb – Ptiss) • τ−1 dt, where Pamb is ambient N2 pressure and τ is a time constant. The probability of DCS [P(DCS)] was predicted from the risk function: P(DCS) = 1−e−r, where r = ∫(PtissN2 − Thr − Pamb) • Pamb–1 dt, and Thr is a threshold parameter. An equation that scaled τ with body mass included a constant (c) and an allometric scaling parameter (n), and the best model included n, Thr, and two c. The final model provided accurate predictions for 58 out of 61 dive profiles for pig, hamster, rat, and mouse. Thus, body mass helped improve the prediction of DCS risk in four mammalian species over a body mass range covering 3 orders of magnitude.

The Marine Mammal Center (TMMC) cares for malnourished California sea lion (CSL) (Zalophus californianus) pups and yearlings every year. Hypoglycemia is a common consequence of malnutrition in young CSLs. Administering dextrose during a hypoglycemic crisis is vital to recovery. Traditional veterinary approaches to treat hypoglycemia pose therapeutic challenges in otariids, as vascular access and catheter maintenance can be difficult. The current approach to a hypoglycemic episode at TMMC is to administer dextrose intravenously (i.v.) by medically trained personnel. Intraperitoneal (i.p.) dextrose administration is an attractive alternative to i.v. administration because volunteer staff with basic training can administer treatment instead of waiting for trained staff to treat. This study compares the effects of i.v., i.p., and no dextrose administration on serum glucose and insulin in clinically healthy, euglycemic CSL yearlings. Three groups of animals, consisting of five sea lions each, were treated with 500 mg/kg dextrose using one of the following routes: i.v., i.p., or no dextrose (control). A jugular catheter was placed, and blood samples were collected at times 0, 5, 15, 30, 60, 120, 180, and 240 min after dextrose administration. I.v. dextrose administration resulted in an increase of serum glucose concentrations from a baseline level of approximately 150 mg/dl to a peak of approximately 350 mg/dl. The resulting hyperglycemia persisted for approximately 2 hr and was associated with an attenuated plasma insulin response compared with most terrestrial mammals. Intraperitoneal dextrose administration resulted in increases of serum glucose to approximately 200 mg/dl, which gradually declined to baseline by 2 hr after dextrose administration. These data suggest that the initial treatment of a hypoglycemic crisis in young malnourished CSLs can be accomplished with i.p. dextrose, thus enabling minimally trained volunteer staff to respond immediately to a crisis. Further studies are needed to determine the most appropriate long-term treatment.

Recent studies of stranded marine mammals indicate that exposure to underwater military sonar may induce pathophysiological responses consistent with decompression sickness (DCS). However, DCS has been difficult to diagnose in marine mammals. We investigated whether blood microparticles (MPs, measured as number/μl plasma), which increase in response to decompression stress in terrestrial mammals, are a suitable biomarker for DCS in marine mammals. We obtained blood samples from trained Steller sea lions ( Eumetopias jubatus, 4 adult females) wearing time-depth recorders that dove to predetermined depths (either 5 or 50 meters). We hypothesized that MPs would be positively related to decompression stress (depth and duration underwater). We also tested the effect of feeding and exercise in isolation on MPs using the same blood sampling protocol. We found that feeding and exercise had no effect on blood MP levels, but that diving caused MPs to increase. However, blood MP levels did not correlate with diving depth, relative time underwater, and presumed decompression stress, possibly indicating acclimation following repeated exposure to depth.

Decompression sickness (DCS; ‘the bends’) is a disease associated with gas uptake at pressure. The basic pathology and cause are relatively well known to human divers. Breath-hold diving marine mammals were thought to be relatively immune to DCS owing to multiple anatomical, physiological and behavioural adaptations that reduce nitrogen gas (N2) loading during dives. However, recent observations have shown that gas bubbles may form and tissue injury may occur in marine mammals under certain circumstances. Gas kinetic models based on measured time-depth profiles further suggest the potential occurrence of high blood and tissue N2 tensions. We review evidence for gas-bubble incidence in marine mammal tissues and discuss the theory behind gas loading and bubble formation. We suggest that diving mammals vary their physiological responses according to multiple stressors, and that the perspective on marine mammal diving physiology should change from simply minimizing N2 loading to management of the N2 load. This suggests several avenues for further study, ranging from the effects of gas bubbles at molecular, cellular and organ function levels, to comparative studies relating the presence/absence of gas bubbles to diving behaviour. Technological advances in imaging and remote instrumentation are likely to advance this field in coming years. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Bubbles in supersaturated tissues and blood occur in beaked whales stranded near sonar exercises, and post-mortem in dolphins bycaught at depth and then hauled to the surface. To evaluate live dolphins for bubbles, liver, kidneys, eyes and blubber–muscle interface of live-stranded and capture-release dolphins were scanned with B-mode ultrasound. Gas was identified in kidneys of 21 of 22 live-stranded dolphins and in the hepatic portal vasculature of 2 of 22. Nine then died or were euthanized and bubble presence corroborated by computer tomography and necropsy, 13 were released of which all but two did not re-strand. Bubbles were not detected in 20 live wild dolphins examined during health assessments in shallow water. Off-gassing of supersaturated blood and tissues was the most probable origin for the gas bubbles. In contrast to marine mammals repeatedly diving in the wild, stranded animals are unable to recompress by diving, and thus may retain bubbles. Since the majority of beached dolphins released did not re-strand it also suggests that minor bubble formation is tolerated and will not lead to clinically significant decompression sickness.

Hydrostatic lung compression in diving marine mammals, with collapsing alveoli blocking gas exchange at depth, has been the main theoretical basis for limiting N2uptake and avoiding gas emboli (GE) as they ascend. However, studies of beached and bycaught cetaceans and sea turtles imply that air-breathing marine vertebrates may, under unusual circumstances, develop GE that result in decompression sickness (DCS) symptoms. Theoretical modelling of tissue and blood gas dynamics of breath-hold divers suggests that changes in perfusion and blood flow distribution may also play a significant role. The results from the modelling work suggest that our current understanding of diving physiology in many species is poor, as the models predict blood and tissue N2levels that would result in severe DCS symptoms (chokes, paralysis and death) in a large fraction of natural dive profiles. In this review, we combine published results from marine mammals and turtles to propose alternative mechanisms for how marine vertebrates control gas exchange in the lung, through management of the pulmonary distribution of alveolar ventilation () and cardiac output/lung perfusion (), varying the level ofin different regions of the lung. Man-made disturbances, causing stress, could alter themismatch level in the lung, resulting in an abnormally elevated uptake of N2, increasing the risk for GE. Our hypothesis provides avenues for new areas of research, offers an explanation for how sonar exposure may alter physiology causing GE and provides a new mechanism for how air-breathing marine vertebrates usually avoid the diving-related problems observed in human divers.

We developed a mathematical model to investigate the effect of lung compression and collapse (pulmonary shunt) on the uptake and removal of O(2), CO(2) and N(2) in blood and tissue of breath-hold diving mammals. We investigated the consequences of pressure (diving depth) and respiratory volume on pulmonary shunt and gas exchange as pressure compressed the alveoli. The model showed good agreement with previous studies of measured arterial O(2) tensions (Pa(O)(2)) from freely diving Weddell seals and measured arterial and venous N(2) tensions from captive elephant seals compressed in a hyperbaric chamber. Pulmonary compression resulted in a rapid spike in Pa(O)(2) and arterial CO(2) tension, followed by cyclical variation with a periodicity determined by Q(tot). The model showed that changes in diving lung volume are an efficient behavioural means to adjust the extent of gas exchange with depth. Differing models of lung compression and collapse depth caused major differences in blood and tissue N(2) estimates. Our integrated modelling approach contradicted predictions from simple models, and emphasised the complex nature of physiological interactions between circulation, lung compression and gas exchange. Overall, our work suggests the need for caution in interpretation of previous model results based on assumed collapse depths and all-or-nothing lung collapse models.

SUMMARY The metabolic costs of foraging and the management of O2 and CO2 stores during breath-hold diving was investigated in three female Steller sea lions (Eumetopias jubatus) trained to dive between 10 and 50 m (N=1142 dives). Each trial consisted of two to eight dives separated by surface intervals that were determined by the sea lion(spontaneous trials) or by the researcher (conditioned trials). During conditioned trials, surface intervals were long enough for O2 to return to pre-dive levels between each dive. The metabolic cost of each dive event (dive+surface interval; DMR) was measured using flow-through respirometry. The respiratory exchange ratio(V̇O2/V̇CO2)was significantly lower during spontaneous trials compared with conditioned trials. DMR was significantly higher during spontaneous trials and decreased exponentially with dive duration. A similar decrease in DMR was not as evident during conditioned trials. DMR could not be accurately estimated from the surface interval (SI) following individual dives that had short SIs (50 s). DMR decreased by 15%, but did not differ significantly from surface metabolic rates (MRS) when dive duration increased from 1 to 7 min. Overall,these data suggest that DMR is almost the same as MRS, and that Steller sea lions incur an O2 debt during spontaneous diving that is not repaid until the end of the dive bout. This has important consequences in differentiating between the actual and `apparent' metabolic rate during diving, and may explain some of the differences in metabolic rates reported in pinniped species.

Changes in buoyancy due to seasonal or abnormal changes in body composition are thought to significantly affect the energy budget of marine mammals through changes in diving costs. We assessed how changes in body composition might alter the foraging efficiency of Steller sea lions Eumetopias jubatus by artificially adjusting the buoyancy of trained individuals. PVC tubes were attached to harnesses worn by Steller sea lions that had been trained to feed at fixed depths (10 to 30 m) and to resurface inside a metabolic dome. Buoyancy was altered to simulate the naturally occurring differences in body composition reported in adult females (~12 to 26% subcutaneous fat). Diving characteristics (transit times and time at depth) and aerobic energy expenditure (gas exchange) were measured. We found that foraging cost decreased with the duration of the dive and increased with dive depth. However, changes in body composition did not affect the diving metabolic rate of Steller sea lions for dives between 10 and 30 m. We propose that Steller sea lions may adjust their diving lung volume to compensate for changes in buoyancy to avoid additional metabolic costs.

Marine mammals have very compliant alveoli and stiff upper airways; an adaptation that allows air to move from the alveoli into the upper airways, during breath-hold diving. Alveolar collapse is thought occur between 30 and 100 m and studies that have attempted to estimate gas exchange at depth have used the simplifying assumption that gas exchange ceases abruptly at the alveolar collapse depth. Here we develop a mathematical model that uses compliance values for the alveoli and upper airspaces, estimated from the literature, to predict volumes of the respiratory system at depth. Any compressibility of the upper airways decreases the volume to contain alveolar air yielding lung collapse pressures 2× that calculated assuming an incompressible upper airway. A simple relationship with alveolar volume was used to predict relative pulmonary shunt at depth. The results from our model agree with empirical data on gas absorption at depth as well as the degree of tracheal compression in forced and free diving mammals.

Three Steller sea lions Eumetopias jubatus were trained to participate in free-swim- ming, open-ocean experiments designed to determine if activity can be used to estimate the ener- getic cost of finding prey at depth. Sea lions were trained to dive to fixed depths of 10 to 50 m, and to re-surface inside a floating dome to measure energy expenditure via gas exchange. A 3-axis accelerometer was attached to the sea lions during foraging. Acceleration data were used to deter- mine the overall dynamic body acceleration (ODBA), a proxy for activity. Results showed that ODBA correlated well with the diving metabolic rate (dive + surface interval) and that the variability in the relationship (r 2 = 0.47, linear regression including Sea lion as a random factor) was similar to that reported for other studies that used heart rate to estimate metabolic rate for sea lions swimming underwater in a 2 m deep water channel. A multivariate analysis suggested that both ODBA and dive duration were important for predicting diving metabolic cost, but ODBA alone predicted foraging cost to within 7% between animals. Consequently, collecting 3-dimensional acceleration data is a simple technique to estimate field metabolic rate of wild Steller sea lions and other diving mammals and birds.

This research investigated whether decompression sickness (DCS) risk or severity could be reduced using drug interventions that are easier to implement and equal to or more efficacious than recompression therapy.Using a rat model of DCS, anti-inflammatory or anticoagulant drugs, including lidocaine, aspirin (ASA), methylprednisolone (MP), alpha-phenyl-N-butylnitrone (PBN), and transsodium crocetinate (TSC) were tested to determine their effect on incidence of DCS, death, and time of symptom onset. Each treatment group consisted of approximately 40 animals that received the drug and approximately 40 controls. Animals were exposed to one of five compression and decompression profiles with pressure ranging from 6.3 ATA (175 fsw) to 8.0 ATA (231 fsw); bottom time was either 60 or 90 min; and decompression rate was either 1.8 or 15 ATA x min(-1). Following decompression, the rats were observed for 30 min while walking on a wheel. DCS was defined as an ambulatory deficit or abnormal breathing.None of the drugs reached statistical significance for all DCS manifestations. Lidocaine post-dive and MP were the only treatments with marginally (P0.15) significant differences in DCS outcomes compared to controls. Lidocaine post-dive significantly decreased the incidence of neurological DCS from 73-51%. MP significantly extended the time of onset of death from DCS from 5.4 min to 7.1 min.Of the treatments investigated, lidocaine given post-dive has the best chance of success in adjuvant therapy of DCS. Future studies might investigate adjuvant drugs given in combination or during recompression.

International audience; Penguins are known to have high pedestrian locomotory costs in comparison to other cursorial birds, but the ecological consequences of this difference have received limited attention. Here we present a method for the accurate estimation of onshore energetics based on measurements of body mass, simple morphometrics and distance moved. The method is shown to be similarly accurate to other field-based estimates of energy expenditure, but has the advantage of logistical simplicity. King penguins spend 30-50% of their time ashore and may walk distances of several kilometres to and from their breeding colonies. However, in such cases the total energetic cost of pedestrian locomotion is estimated to be only 1.0% of the energy expended whilst ashore. Thus, despite a high instantaneous cost, pedestrian locomotion is a small and possibly negligible component of total energy turnover in king penguins.

King penguins (Aptenodytes patagonicus) are one of the greatest consumers of marine resources. However, while their influence on the marine ecosystem is likely to be significant, only an accurate knowledge of their energy demands will indicate their true food requirements. Energy consumption has been estimated for many marine species using the heart rate–rate of oxygen consumption ( f H – V ˙ O 2 ) technique, and the technique has been applied successfully to answer eco-physiological questions. However, previous studies on the energetics of king penguins, based on developing or applying this technique, have raised a number of issues about the degree of validity of the technique for this species. These include the predictive validity of the present f H – V ˙ O 2 equations across different seasons and individuals and during different modes of locomotion. In many cases, these issues also apply to other species for which the f H – V ˙ O 2 technique has been applied. In the present study, the accuracy of three prediction equations for king penguins was investigated based on validity studies and on estimates of V ˙ O 2 from published, field fH data. The major conclusions from the present study are: (1) in contrast to that for walking, the f H – V ˙ O 2 relationship for swimming king penguins is not affected by body mass; (2) prediction equation (1), log ( V ˙ O 2 ) = - 0.279 + 1.24 log ( f H ) + 0.0237 t - 0.0157 log ( f H ) t , derived in a previous study, is the most suitable equation presently available for estimating V ˙ O 2 in king penguins for all locomotory and nutritional states. A number of possible problems associated with producing an f H – V ˙ O 2 relationship are discussed in the present study. Finally, a statistical method to include easy-to-measure morphometric characteristics, which may improve the accuracy of f H – V ˙ O 2 prediction equations, is explained.

We measured esophageal pressures, respiratory flow rates, and expired O 2 and CO 2 in six adult bottlenose dolphins ( Tursiops truncatus ) during voluntary breaths and maximal (chuff) respiratory efforts. The data were used to estimate the dynamic specific lung compliance (s C L ), the O 2 consumption rate ( V O 2 ) and CO 2 production rates ( V CO 2 ) during rest. Our results indicate that bottlenose dolphins have the capacity to generate respiratory flow rates that exceed 130 l s −1 and 30 l s −1 during expiration and inspiration, respectively. The esophageal pressures indicated that expiration is passive during voluntary breaths, but active during maximal efforts, whereas inspiration is active for all breaths. The average s C L of dolphins was 0.31±0.04 cmH 2 O −1 , which is considerably higher than that of humans (0.08 cmH 2 O −1 ) and that previously measured in a pilot whale (0.13 cmH 2 O −1 ). The average estimated V O 2 and V CO 2 using our breath-by-breath respirometry system ranged from 0.857 to 1.185 l O 2 min −1 and 0.589 to 0.851 l CO 2 min −1 , respectively, which is similar to previously published metabolic measurements from the same animals using conventional flow-through respirometry. In addition, our custom-made system allows us to approximate end tidal gas composition. Our measurements provide novel data for respiratory physiology in cetaceans, which may be important for clinical medicine and conservation efforts.

Heart-rate (fH) changes during diving and exercise are well documented for marine mammals, but changes in stroke volume (SV) and cardiac output (CO) are much less known. We hypothesized that both SV and CO are also modified following intense exercise. Using transthoracic ultrasound Doppler at the level of the aortic valve, we compared blood flow velocities in the left ventricle and cardiac frequencies during rest and at 1, 3 and 4 min after a bout of exercise in 13 adult bottlenose dolphins (Tursiops truncatus, six male and seven female, body mass range: 143-212 kg). Aortic cross sectional area and ventricle blood velocity at the aortic valve were used to calculate SV, which together with fH, provided estimates of left CO at rest and following exercise. The fH and SV stabilized approximately 4-7 sec following the post-respiratory tachycardia, so only data after the fH had stabilized were used for analysis and comparison. There were significant increases in fH, SV, and CO associated with each breath. At rest, fH, SV, and CO were uncorrelated with body mass, and averaged 41±8 beats min−1, 136±19 ml, and 5513±1182 l min−1, respectively. One minute following high intensity exercise, the cardiac variables had increased by 104±43%, 63±11%, and 234±84%, respectively. All variables remained significantly elevated in all animals for at least 4 min after the exercise. These baseline values provide the first data on stroke volume and cardiac output in awake and unrestrained cetaceans in water.

A mathematical model was created that predicted blood and tissue N(2) tension (P(N2)) during breath-hold diving. Measured muscle P(N2) from the bottlenose dolphin after diving repeatedly to 100 m (Tursiops truncatus [Ridgway and Howard, 1979, Science, 4423, 1182-1183]) was compared with predictions from the model. Lung collapse was modelled as a 100% pulmonary shunt which yielded tissue P(N2) similar to those reported for the dolphin. On the other hand, predicted muscle P(N2) for an animal with a dive response, reducing cardiac output by 66% from surface values (20.5 to 6.8l x min(-1)), also agreed well with observed values in the absence of lung collapse. In fact, modelling indicated that both cardiovascular adjustments and dive behaviour are important in reducing N2 uptake during diving and enhancing safe transfer of tissue and blood N2 back to the lung immediately before coming to the surface. In particular, diving bradycardia during the descent and bottom phase together with a reduced ascent rate and increase in heart rate reduced mixed venous P(N2) upon return to the surface by as much as 45%. This has important implications as small reductions in inert gas load (approximately 5%) can substantially reduce decompression sickness (DCS) risk by as much as 50% (Fahlman et al., 2001, J. Appl. Physiol. 91, 2720-2729).

6 PAGES; International audience; We describe a method that allows prediction of resting metabolic rate (RMR, ml O2 Æ min1) in adult male and female king penguins on shore by measuring body mass (Mb) and the length of the foot, flipper and beak. This method is accurate, underestimating measured RMR (n=114) by 4% in a data set consisting of 44 birds (33 males and 11 females). Measurement error was unbiased with respect to fasting duration and can therefore estimate RMR during any stage of fasting. This new method provides significant cost and logistical savings when estimating RMR during fieldwork, allowing RMR of a large number of birds to be measured quickly. These findings suggest the possibility that the use of Mb and morphometrics will allow development of general and specific equations to estimate RMR in other species.

The acute Hypoxic Ventilatory Response (HVR) is an important component of human hypoxia tolerance, hence presumably physiological adaptation to high altitude. We measured the isocapnic HVR (L min(-1) %(-1)) in two genetically divergent low altitude southern African populations. The HVR does not differ between African Xhosas (X) and Caucasians (C) (X:-0.34+/-0.36; C:-0.42+/-0.33; P0.34), but breathing patterns do. Among all Xhosa subjects, size-independent tidal volume was smaller (X: 0.75+/-0.20; C: 1.11+/-0.32 L; P0.01), breathing frequency higher (X: 22.2+/-5.7; C: 14.3+/-4.2 breaths min(-1); P0.01) and hypoxic oxygen saturation lower than among Caucasians (X: 78.4+/-4.7%; C: 81.7+/-4.7%; P0.05). The results remained significant if subjects from Xhosa and Caucasian groups were matched for gender, body mass index and menstrual cycle phase in the case of females. The latter also employed distinct breathing patterns between populations in normoxia. High repeatability (intra-class correlation coefficient) of the HVR in both populations (0.77-0.87) demonstrates that one of the prerequisites for natural selection, consistent between-individual variation, is met. Finally, we explore possible relationships between inter-population genetic distances and HVR differences among Xhosa, European, Aymara Amerindians, Tibetan and Chinese populations. Inter-population differences in the HVR are not attributable to genetic distance (Mantel Z-test, P = 0.59). The results of this study add novel support for the hypothesis that differences in the HVR, should they be found between other human populations, may reflect adaptation to hypoxia rather than genetic divergence through time.

SUMMARYThe main objective of this study was to determine heart rate(fh) and the energetic costs of specific behaviours of king penguins while ashore and while foraging at sea during their breeding period. In particular, an estimate was made of the energetic cost of diving in order to determine the proportion of dives that may exceed the calculated aerobic dive limit (cADL; estimated usable O2 stores/estimated rate of oxygen consumption during diving).An implanted data logger enabled fh and diving behaviour to be monitored from 10 free-ranging king penguins during their breeding period. Using previously determined calibration equations, it was possible to estimate rate of oxygen consumption(V̇O2) when the birds were ashore and during various phases of their foraging trips. Diving behaviour showed a clear diurnal pattern, with a mixture of deep (>40 m),long (>3 min) and shallow (Rates of oxygen consumption estimated from these (and other) values of fh indicate that when at sea, metabolic rate (MR) was 83%greater than that when the birds were ashore [3.15 W kg–1(–0.71, +0.93), where the values in parentheses are the computed standard errors of the estimate], while during diving bouts and dive cycles,it was 73% greater than the `ashore' value. Although estimated MR during the total period between dive bouts was not significantly different from that during dive bouts [5.44 W kg–1 (–0.30, +0.32)], MR during the first hour following a dive bout was 52% greater than that during a diving bout. It is suggested that this large increase following diving(foraging) activity is, at least in part, the result of rewarming the body,which occurs at the end of a diving bout. From the measured behaviour and estimated values of V̇O2, it was evident that approximately 35% of the dives were in excess of the cADL. Even if V̇O2 during diving was assumed to be the same as when the birds were resting on water,approximately 20% of dives would exceed the cADL. As V̇O2 during diving is, in fact, that estimated for a complete dive cycle, it is quite feasible that V̇O2 during diving itself is less than that measured for birds resting in water. It is suggested that the regional hypothermia that has been recorded in this species during diving bouts may be at least a contributing factor to such hypometabolism.

King penguins ( Aptenodytes patagonicus) may fast for up to 30 days during their breeding period. As such extended fasting may affect the relationship between the rate of O2consumption (V̇o2) and heart rate ( fH), five male king penguins were exercised at various speeds on repeated occasions during a fasting period of 24–31 days. In addition, V̇o2and fHwere measured in the same animals during rest in cold air and water (4°C). V̇o2and fHat rest and V̇o2during exercise decreased with fasting. There was a significant relation between V̇o2and fH( r2= 0.56) that was improved by including speed, body mass ( Mb), number of days fasting ( t), and a cross term between fHand t ( r2= 0.92). It was concluded that there was a significant change in the V̇o2- fHrelationship with fasting during exercise. As t is measurable in the field and was shown to be significant and, therefore, a practical covariate, a regression equation for use when birds are ashore was obtained by removing speed and Mb. When this equation was used, predicted V̇o2was in good agreement with the observed data, with an overall error of 3.0%. There was no change in the V̇o2- fHrelationship in penguins at rest in water.

We modelled the kinetics of H2 flux during gas uptake and elimination in conscious pigs exposed to hyperbaric H2. The model used a physiological description of gas flux fitted to the observed decompression sickness (DCS) incidence in two groups of pigs: untreated controls, and animals that had received intestinal injections of H2-metabolizing microbes that biochemically eliminated some of the H2 stored in the pigs’ tissues. To analyse H2 flux during gas uptake, animals were compressed in a dry chamber to 24 atm (ca 88% H2, 9% He, 2% O2, 1% N2) for 30–1440 min and decompressed at 0.9 atm min −1 (n=70) . To analyse H2 flux during gas elimination, animals were compressed to 24 atm for 3 h and decompressed at 0.45–1.8 atm min−1(n=58). Animals were closely monitored for 1 h post-decompression for signs of DCS. Probabilistic modelling was used to estimate that the exponential time constant during H2 uptake (τin) and H2 elimination (τout) were 79±25 min and 0.76±0.14 min, respectively. Thus, the gas kinetics affecting DCS risk appeared to be substantially faster for elimination than uptake, which is contrary to customary assumptions of gas uptake and elimination kinetic symmetry. We discuss the possible reasons for this asymmetry, and why absolute values of H2 kinetics cannot be obtained with this approach.

SUMMARYHumboldt penguins Spheniscus humboldti in captivity and free-living Magellanic penguins S. magellanicus were fitted with loggers to determine beak angles during breathing. The Humboldt penguins were also fitted with masks for determining rates of air flow during breathing. During periods of higher gas exchange requirement, Humboldt penguins opened their beaks during inspiration, where tidal volume was linearly correlated with both change in beak angle and maximum beak angle, closed them slightly during the final stages of inspiration and finally closed them during expiration. Substantial differences were apparent between individuals. Contrary to the condition proposed for most birds, our data suggest that expiration is passive during periods of high respiratory tidal volumes, and that the increased resistance of the respiratory pathway serves to slow air flow so as to maximize gas exchange in the lungs. During foraging, Magellanic penguins at the surface between dives showed similar breathing patterns but maximum beak angles were much higher and breath cycle time shorter, as would be expected for animals attempting to maximize gas exchange. Both maximum beak angle per breath and breath frequency changed systematically over the surface pause; both were initially high, then decreased to a low before rising again to a maximum just before diving. Based on known changes in tidal volume with beak angle derived from Humboldt penguins, a simple model is proposed to examine rates of gas exchange over the surface pause. This indicates that the observed patterns do not maximize the rate of transfer of oxygen over the whole of the surface pause but are rather concerned with an initial rapid accumulation of oxygen in the tissues followed by effective carbon dioxide release.

Attaching bio-telemetry or -logging devices ('tags') to marine animals for research and monitoring adds drag to streamlined bodies, affecting posture, swimming gaits and energy balance. These costs have never been measured in free-swimming cetaceans. To examine the effect of drag from a tag on metabolic rate, cost of transport, and swimming behavior, four captive male dolphins (Tursiops truncatus) were trained to swim a set course, either non-instrumented (n = 7) or instrumented with a tag (DTAG2; n = 12), and surface exclusively in a flow-through respirometer where oxygen consumption (V̇O2) and carbon dioxide production (V̇CO2; mL kg-1 min-1) rates were measured and respiratory exchange ratio (V̇O2/V̇CO2) was calculated. Tags did not significantly affect individual mass-specific oxygen consumption, Physical Activity Ratios (exercise V̇O2/resting V̇O2), total or net cost of transport (COT, J m-1 kg-1) or locomotor costs during swimming or two-minute recovery phases. However, individuals swam significantly slower when tagged (by ~11%; mean±s.d. 3.31±0.35 m s-1) compared to when non-instrumented (3.73±0.41 m s-1). A combined theoretical and Computational Fluid Dynamics (CFD) model estimating drag forces and power exertion during swimming suggests drag loading and energy consumption are reduced at lower swimming speeds. Bottlenose dolphins in the specific swimming task in this experiment slowed to the point where the tag yielded no increases in drag or power, while showing no difference in metabolic parameters when instrumented with a DTAG2. These results, and our observations, suggest that animals modify their behavior to maintain metabolic output and energy expenditure when faced with tag-induced drag.

Northern elephant seals (Mirounga angustirostris) (NES) are known to be deep, long-duration divers and to sustain long-repeated patterns of breath-hold, or apnea. Some phocid dives remain within the bounds of aerobic metabolism, accompanied by physiological responses inducing lung compression, bradycardia, and peripheral vasoconstriction. Current data suggest an absence of type IIb fibers in pinniped locomotory musculature. To date, no fiber type data exist for NES, a consummate deep diver. In this study, NES were biopsied in the wild. Ontogenetic changes in skeletal muscle were revealed through succinate dehydrogenase (SDH) based fiber typing. Results indicated a predominance of uniformly shaped, large type I fibers and elevated myoglobin (Mb) concentrations in the longissimus dorsi (LD) muscle of adults. No type II muscle fibers were detected in any adult sampled. This was in contrast to the juvenile animals that demonstrated type II myosin in Western Blot analysis, indicative of an ontogenetic change in skeletal muscle with maturation. These data support previous hypotheses that the absence of type II fibers indicates reliance on aerobic metabolism during dives, as well as a depressed metabolic rate and low energy locomotion. We also suggest that the lack of type IIb fibers (adults) may provide a protection against ischemia reperfusion (IR) injury in vasoconstricted peripheral skeletal muscle.

Recent cetacean mass strandings in close temporal and spatial association with sonar activity has raised the concern that anthropogenic sound may harm breath-hold diving marine mammals. Necropsy results of the stranded whales have shown evidence of bubbles in the tissues, similar to those in human divers suffering from decompression sickness (DCS). It has been proposed that changes in behavior or physiological responses during diving could increase tissue and blood N2 levels, thereby increasing DCS risk. Dive data recorded from sperm, killer, long-finned pilot, Blainville's beaked and Cuvier's beaked whales before and during exposure to low- (1–2 kHz) and mid- (2–7 kHz) frequency active sonar were used to estimate the changes in blood and tissue N2 tension (PN2). Our objectives were to determine if differences in (1) dive behavior or (2) physiological responses to sonar are plausible risk factors for bubble formation. The theoretical estimates indicate that all species may experience high N2 levels. However, unexpectedly, deep diving generally result in higher end-dive PN2 as compared with shallow diving. In this focused review we focus on three possible explanations: (1) We revisit an old hypothesis that CO2, because of its much higher diffusivity, forms bubble precursors that continue to grow in N2 supersaturated tissues. Such a mechanism would be less dependent on the alveolar collapse depth but affected by elevated levels of CO2 following a burst of activity during sonar exposure. (2) During deep dives, a greater duration of time might be spent at depths where gas exchange continues as compared with shallow dives. The resulting elevated levels of N2 in deep diving whales might also make them more susceptible to anthropogenic disturbances. (3) Extended duration of dives even at depths beyond where the alveoli collapse could result in slow continuous accumulation of N2 in the adipose tissues that eventually becomes a liability. Publisher PDF