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Understanding the behavioural ecology of endangered taxa can inform conservation strategies. The activity budgets of the loggerhead turtle Caretta caretta are still poorly understood because many tracking methods show only horizontal displacement and ignore dives and associated behaviours. However, time-depth recorders have enabled researchers to identify flat, U-shaped dives (or type 1a dives) and these are conventionally labelled as resting dives on the seabed because they involve no vertical displacement of the animal. Video- and acceleration-based studies have demonstrated this is not always true. Focusing on sea turtles nesting on the Cabo Verde archipelago, we describe a new metric derived from magnetometer data, absolute angular velocity, that integrates indices of angular rotation in the horizontal plane to infer activity. Using this metric, we evaluated the variation in putative resting behaviours during the bottom phase of type 1a dives for 5 individuals over 13 to 17 d at sea during a single inter-nesting interval (over 75 turtle d in total). We defined absolute resting within the bottom phase of type 1a dives as periods with no discernible acceleration or angular movement. Whilst absolute resting constituted a significant proportion of each turtle’s time budget for this 1a dive type, turtles allocated 16−38% of their bottom time to activity, with many dives being episodic, comprised of intermittent bouts of rest and rotational activity. This implies that previously considered resting behaviours are complex and need to be accounted for in energy budgets, particularly since energy budgets may impact conservation strategies. © The authors 2021. Open Access under Creative Commons by Attribution Licence. Use, distribution and reproduction are unrestricted. Authors and original publication must be credited

This work was funded by the Marine Mammal Commission (MMC19-173). The Office of Naval Research funded the bioenergetic workshop (N000142012392) that provided support for this work. Bioenergetic approaches are increasingly used to understand how marine mammal populations could be affected by a changing and disturbed aquatic environment. There remain considerable gaps in our knowledge of marine mammal bioenergetics, which hinder the application of bioenergetic studies to inform policy decisions. We conducted a priority-setting exercise to identify high-priority unanswered questions in marine mammal bioenergetics, with an emphasis on questions relevant to conservation and management. Electronic communication and a virtual workshop were used to solicit and collate potential research questions from the marine mammal bioenergetic community. From a final list of 39 questions, 11 were identified as ‘key’ questions because they received votes from at least 50% of survey participants. Key questions included those related to energy intake (prey landscapes, exposure to human activities) and expenditure (field metabolic rate, exposure to human activities, lactation, time-activity budgets), energy allocation priorities, metrics of body condition and relationships with survival and reproductive success and extrapolation of data from one species to another. Existing tools to address key questions include labelled water, animal-borne sensors, mark-resight data from long-term research programs, environmental DNA and unmanned vehicles. Further validation of existing approaches and development of new methodologies are needed to comprehensively address some key questions, particularly for cetaceans. The identification of these key questions can provide a guiding framework to set research priorities, which ultimately may yield more accurate information to inform policies and better conserve marine mammal populations. Publisher PDF

Estimates of the energetic costs of locomotion (COL) at different activity levels are necessary to answer fundamental eco-physiological questions and to understand the impacts of anthropogenic disturbance to marine mammals. We combined estimates of energetic costs derived from breath-by-breath respirometry with measurements of overall dynamic body acceleration (ODBA) from biologging tags to validate ODBA as a proxy for COL in trained common bottlenose dolphins (Tursiops truncatus). We measured resting metabolic rate (RMR); mean individual RMR was 0.71–1.42 times that of a similarly sized terrestrial mammal and agreed with past measurements that used breath-by-breath and flow-through respirometry. We also measured energy expenditure during submerged swim trials, at primarily moderate exercise levels. We subtracted RMR to obtain COL, and normalized COL by body size to incorporate individual swimming efficiencies. We found both mass-specific energy expenditure and mass-specific COL were linearly related with ODBA. Measurements of activity level and cost of transport (the energy required to move a given distance) improve understanding of the COL in marine mammals. The strength of the correlation between ODBA and COL varied among individuals, but the overall relationship can be used at a broad scale to estimate the energetic costs of disturbance and daily locomotion costs to build energy budgets, and investigate the costs of diving in free-ranging animals where bio-logging data are available. We propose that a similar approach could be applied to other cetacean species.

Pulmonary function testing was performed in 3 bottlenose dolphins Tursiops truncatus (1 female and 2 males) under managed care during a 2 yr period to assess whether these data provide diagnostic information about respiratory health. Pulmonary radiographs and standard clinical testing were used to evaluate the pulmonary health of each dolphin. The female dolphin (F1) had evidence of chronic pulmonary fibrosis, and 1 male (M2) developed pneumonia during the study. Pulmonary function data were collected from maximal respiratory efforts in water and from spontaneous breaths while beached. From these data, the flow-volume relationship, the flow measured between 25 and 75% of the expired vital capacity (mid forced expiratory flow, FEF25%-75%), and the percent of the vital capacity (VC) at the peak expiratory flow (%VCPEF), were evaluated and compared with the diagnostic assessment. For maximal respiratory manoeuvres in water, there were no differences in FEF25%-75% or %VCPEF, and the flow-volume relationship showed a consistent pattern for F1. Additionally, FEF25%-75% and %VCPEF decreased by 27 and 52%, respectively, and the flow-volume relationship showed clear flow limitations with emerging disease in M2. While spontaneously breathing on land, M2 also showed a 49% decrease in %VCPEF and changes in the flow-volume relationship, indicating flow limitations following the development of pneumonia. Based on these preliminary results, we suggest that pulmonary function testing should be given more attention as a non-invasive and possibly adjunctive diagnostic tool to evaluate lung health of dolphins under managed care and in the wild.

Plasticity in the cardiac function of a marine mammal facilitates rapid adjustments to the contrasting metabolic demands of breathing at the surface and diving during an extended apnea. By matching their heart rate ( f H ) to their immediate physiological needs, a marine mammal can improve its metabolic efficiency and maximize the proportion of time spent underwater. Respiratory sinus arrhythmia (RSA) is a known modulation of f H that is driven by respiration and has been suggested to increase cardiorespiratory efficiency. To investigate the presence of RSA in cetaceans and the relationship between f H , breathing rate ( f R ) and body mass ( M b ), we measured simultaneous f H and f R in five cetacean species in human care. We found that a higher f R was associated with a higher mean instantaneous f H (i f H ) and minimum i f H of the RSA. By contrast, f H scaled inversely with M b such that larger animals had lower mean and minimum i f H s of the RSA. There was a significant allometric relationship between maximum i f H of the RSA and M b , but not f R , which may indicate that this parameter is set by physical laws and not adjusted dynamically with physiological needs. RSA was significantly affected by f R and was greatly reduced with small increases in f R . Ultimately, these data show that surface f H s of cetaceans are complex and the f H patterns we observed are controlled by several factors. We suggest the importance of considering RSA when interpreting f H measurements and particularly how f R may drive f H changes that are important for efficient gas exchange. This article is part of the theme issue ‘Measuring physiology in free-living animals (Part I)’.

Walruses rely on sea-ice to efficiently forage and rest between diving bouts while maintaining proximity to prime foraging habitat. Recent declines in summer sea ice have resulted in walruses hauling out on land where they have to travel farther to access productive benthic habitat while potentially increasing energetic costs. Despite the need to better understand the impact of sea ice loss on energy expenditure, knowledge about metabolic demands of specific behaviours in walruses is scarce. In the present study, 3 adult female Pacific walruses (Odobenus rosmarus divergens) housed in professional care participated in flow-through respirometry trials to measure metabolic rates while floating inactive at the water surface during a minimum of 5 min, during a 180 s stationary dive, and while swimming ∼90 m horizontally underwater. Metabolic rates during stationary dives (3.82±0.56 l O2 min−1) were lower than those measured at the water surface (4.64±1.04 l O2 min−1), which did not differ from rates measured during subsurface swimming (4.91±0.77 l O2 min−1). Thus, neither stationary diving nor subsurface swimming resulted in metabolic rates above those exhibited by walruses at the water surface. These results suggest that walruses minimize their energetic investment during underwater behaviours as reported for other marine mammals. Although environmental factors experienced by free-ranging walruses (e.g. winds or currents) likely affect metabolic rates, our results provide important information for understanding how behavioural changes affect energetic costs and can be used to improve bioenergetics models aimed at predicting the metabolic consequences of climate change on walruses.

Lung function (breath duration, respiratory flow [], and tidal volume [VT]), and end-expiratory O2 were measured in 19 adult bottlenose dolphins (Tursiops spp.) while at rest in water or beached for up to 10 min. The results show that inspiratory VT, expiratory VT, or inspiratory did not differ on land or in water. The average expiratory for all dolphins on land decreased by 16%, and the expiratory and total breath durations increased by 5% and 4%, respectively, compared with in water. There were temporal changes observed during beaching, where expired and inspired VT and inspired decreased by 13%, 16%, and 9%, respectively, after 10 min on land. These data suggest that dolphins compensate for the effect of gravity by adjusting respiration to maintain alveolar ventilation and gas exchange, but during extended durations, the increased work of breathing may impede ventilation and gas exchange. Continuous monitoring of lung function and gas exchange may help prevent long-term damage during out-of-water medical procedures, optimize animal transport conditions, and improve survival during stranding events.

Background and objectives Ischemic events, such as ischemic heart disease and stroke, are the number one cause of death globally. Ischemia prevents blood, carrying essential nutrients and oxygen, from reaching tissues, leading to cell and tissue death, and eventual organ failure. While humans are relatively intolerant to ischemic events, other species, such as marine mammals, have evolved a unique tolerance to chronic ischemia/reperfusion during apneic diving. To identify possible molecular features of an increased tolerance for apnea, we examined changes in gene expression in breath-holding dolphins. Methodology Here, we capitalized on the adaptations possesed by bottlenose dolphins (Tursiops truncatus) for diving as a comparative model of ischemic stress and hypoxia tolerance to identify molecular features associated with breath holding. Given that signals in the blood may influence physiological changes during diving, we used RNA-Seq and enzyme assays to examine time-dependent changes in gene expression in the blood of breath-holding dolphins. Results We observed time-dependent upregulation of the arachidonate 5-lipoxygenase (ALOX5) gene and increased lipoxygenase activity during breath holding. ALOX5 has been shown to be activated during hypoxia in rodent models, and its metabolites, leukotrienes, induce vasoconstriction. Conclusions and implications The upregulation of ALOX5 mRNA occurred within the calculated aerobic dive limit of the species, suggesting that ALOX5 may play a role in the dolphin’s physiological response to diving, particularly in a pro-inflammatory response to ischemia and in promoting vasoconstriction. These observations pinpoint a potential molecular mechanism by which dolphins, and perhaps other marine mammals, respond to the prolonged breath holds associated with diving.

Among the many factors that influence the cardiovascular adjustments of marine mammals is the act of respiration at the surface, which facilitates rapid gas exchange and tissue re-perfusion between dives. We measured heart rate (fH) in six adult male bottlenose dolphins (Tursiops truncatus) spontaneously breathing at the surface to quantify the relationship between respiration and fH, and compared this with fH during submerged breath-holds. We found that dolphins exhibit a pronounced respiratory sinus arrhythmia (RSA) during surface breathing, resulting in a rapid increase in fH after a breath followed by a gradual decrease over the following 15–20 s to a steady fH that is maintained until the following breath. RSA resulted in a maximum instantaneous fH (ifH) of 87.4±13.6 beats min−1 and a minimum ifH of 56.8±14.8 beats min−1, and the degree of RSA was positively correlated with the inter-breath interval (IBI). The minimum ifH during 2 min submerged breath-holds where dolphins exhibited submersion bradycardia (36.4±9.0 beats min−1) was lower than the minimum ifH observed during an average IBI; however, during IBIs longer than 30 s, the minimum ifH (38.7±10.6 beats min−1) was not significantly different from that during 2 min breath-holds. These results demonstrate that the fH patterns observed during submerged breath-holds are similar to those resulting from RSA during an extended IBI. Here, we highlight the importance of RSA in influencing fH variability and emphasize the need to understand its relationship to submersion bradycardia.

We analysed 3680 dives from 23 satellite-linked tags deployed on Cuvier's beaked whales to assess the relationship between long duration dives and inter-deep dive intervals and to estimate aerobic dive limit (ADL). The median duration of presumed foraging dives was 59 min and 5% of dives exceeded 77.7 min. We found no relationship between the longest 5% of dive durations and the following inter-deep dive interval nor any relationship with the ventilation period immediately prior to or following a long dive. We suggest that Cuvier's beaked whales have low metabolic rates, high oxygen storage capacities and a high acid-buffering capacity to deal with the by-products of both aerobic and anaerobic metabolism, which enables them to extend dive durations and exploit their bathypelagic foraging habitats.

Lung function (breath duration, respiratory flow [

Among the many factors that influence the cardiovascular adjustments of marine mammals is the act of respiration at the surface, which facilitates rapid gas exchange and tissue re-perfusion between dives. We measured heart rate (

The air volume in the respiratory system of marine tetrapods provides a store of O2 to fuel aerobic metabolism during dives; however, it can also be a liability, as the associated N2 can increase the risk of decompression sickness. In order to more fully understand the physiological limitations of different air-breathing marine vertebrates, it is therefore important to be able to accurately estimate the air volume in the respiratory system during diving. One method that has been used to do so is to calculate the air volume from glide phases – periods of movement during which no thrust is produced by the animal – which many species conduct during ascent periods, when gases are expanding owing to decreasing hydrostatic pressure. This method assumes that there is conservation of mass in the respiratory system, with volume changes only driven by pressure. In this Commentary, we use previously published data to argue that both the respiratory quotient and differences in tissue and blood gas solubility potentially alter the mass balance in the respiratory system throughout a dive. Therefore, near the end of a dive, the measured volume of gas at a given pressure may be 12–50% less than from the start of the dive; the actual difference will depend on the length of the dive, the cardiac output, the pulmonary shunt and the metabolic rate. Novel methods and improved understanding of diving physiology will be required to verify the size of the effects described here and to more accurately estimate the volume of gas inhaled at the start of a dive.

Costs of sound production have been investigated only sparsely in cetaceans, despite recent efforts to understand how increasing anthropogenic noise affects these animals that rely extensively on sound for communication and foraging. Theoretical estimates suggest that metabolic costs of whistling

We measured respiratory flow, breath duration, and calculated tidal volume (VT) in nine belugas (Delphinapterus leucas, mean measured body mass: 628 ± 151 kg, n = 5) housed in managed care facilities. Both spontaneous (resting at station) and trained maximal respirations (chuffs) were measured. The mean (±s.d.) inspiratory VT for spontaneous breaths (16.7 ± 4.7 l, range: 7.5-18.7 l) was larger than those predicted based on respiratory scaling equations from terrestrial mammals and was 32 ± 10% of estimated total lung capacity (TLCest) based on an equation from static measurements made on a range of cetaceans and pinniped lungs, and 52 ± 18% of estimated vital capacities (VC, mean: 27.7 ± 8.9 l, range: 16.7-40.3 l) based on respiratory measurements obtained during trained maximal respirations. Expiratory flow (V˙exp, spontaneous: 26.1 ± 5.5 l s-1, chuff: 66.8 ± 22.5 l s-1) was significantly higher as compared with inspiratory flow (V˙insp, spontaneous: 22.3 ± 4.6 l s-1, chuff: 30.1 ± 8.4 l s-1), and the maximal expiratory flow recorded was 212 l s-1. The breath duration was shorter for forced breaths (Expiration: 518 ± 101 ms; Inspiration: 905 ± 170 ms; Total: 1423 ± 227 ms) as compared with spontaneous breaths (Expiration: 995 ± 176 ms; Inspiration: 1098 ± 219 ms; Total: 2093 ± 302 ms). These data provide baseline estimates of the respiratory capacity of belugas.

In the present study, we examined lung function in healthy resting adult (born in 2003) Pacific walruses (

Toothed whales depend on sound for communication and foraging, making them potentially vulnerable to acoustic masking from increasing anthropogenic noise. Masking effects may be ameliorated by higher amplitudes or rates of calling, but such acoustic compensation mechanisms may incur energetic costs if sound production is expensive. The costs of whistling in bottlenose dolphins (Tursiops truncatus) have been reported to be much higher (20-50% of resting metabolic rate, RMR) than theoretical predictions (0.5-1% of RMR). Here we address this dichotomy by measuring the change in the resting O2 consumption rate (V̇O2), a proxy for RMR, in three post-absorptive bottlenose dolphins during whistling and silent trials, concurrent with simultaneous measurement of acoustic output using a calibrated hydrophone array. The experimental protocol consisted of a two-minute baseline period to establish RMR, followed by a two-minute voluntary resting surface apnea, with or without whistling as cued by the trainers, followed by a five-minute resting period to measure recovery costs. Daily fluctuations in V̇O2 were accounted for by subtracting the baseline RMR from the recovery costs to estimate the cost of apnea with and without whistles relative to RMR. Analysis of 52 sessions containing 1162 whistles showed that whistling did not increase metabolic cost (P>0.1, +4.2±6.9%) (mean±s.e.m.) as compared with control trials (-0.5±5.9%) (mean±s.e.m.). Thus, we reject the hypothesis that whistling is costly for bottlenose dolphins, and conclude that vocal adjustments such as the Lombard response to noise do not represent large direct energetic costs for communicating toothed whales.

Respiratory flow, expired O-2, and CO2 were measured during voluntary participation while spontaneously breathing in 13 confirmed healthy, male California sea lions (Zalophus californianus; body mass [M-b] range: 49 to 130 kg). Expiratory and inspiratory flow ((V) over dot(exp) and (V) over dot(insp)), tidal volume ((V) over dot(Texp), and V-Tinsp), and breath durations (T-exp, T-insp and T-tot) were collected on land (lying down in sternal recumbency and sitting up) and floating in water to test the hypothesis that lung function changes with body position and on land versus in water. For sea lions on land, no differences were seen in any of the lung function values when comparing lying down versus sitting up. However, when comparing animals on land versus in water, both T(exp)( )and T-insp decreased and (V) over dot(exp) and (V) over dot(insp) increased, while the V(T)(exp )and V-T(insp) remained the same. The resting mass-specific VT (25.1 +/- 1.7 ml kg(-1)) in the current study was approximately 24 to 30% of the estimated total lung capacity. We also measured breath-by-breath gas uptake to determine the O-2 consumption rates ((V) over dotO(2)) and CO2 production rates ((V) over dotCO(2)) during rest on land and in water. There were no differences in (V) over dotO(2) or (V) over dotCO(2) on land as compared with water, and the average estimated values were 0.58 +/- 0.22 1 O-2 min(-1) (range: 0.24 to 1.01 1 O-2 min(-1)) and 0.50 +/- 0.19 1 CO2 min(-1) (range: 0.22 to 0.89 1 CO2 min(-1)), respectively, which agrees with results from other studies on otariids. Additionally, the allometric mass-exponent for V-T and (V) over dotO(2) were 1.13 to 1.20 and 0.86, respectively. These data are the first reported estimates of metabolic rate and lung function in confirmed healthy California sea lions.

In the present study, we examined lung function in healthy resting adult (born in 2003) Pacific walruses (Odobenus rosmarus divergens) by measuring respiratory flow ([Formula: see text]) using a custom-made pneumotachometer. Three female walruses (670-1025 kg) voluntarily participated in spirometry trials while spontaneously breathing on land (sitting and lying down in sternal recumbency) and floating in water. While sitting, two walruses performed active respiratory efforts, and one animal participated in lung compliance measurements. For spontaneous breaths, [Formula: see text] was lower when walruses were lying down (e.g. expiration: 7.1±1.2 l s-1) as compared with in water (9.9±1.4 l s-1), while tidal volume (VT, 11.5±4.6 l), breath duration (4.6±1.4 s) and respiratory frequency (7.6±2.2 breaths min-1) remained the same. The measured VT and specific dynamic lung compliance (0.32±0.07 cmH2O-1) for spontaneous breaths were higher than those estimated for similarly sized terrestrial mammals. VT increased with body mass (allometric mass-exponent=1.29) and ranged from 3% to 43% of the estimated total lung capacity (TLCest) for spontaneous breaths. When normalized for TLCest, the maximal expiratory [Formula: see text] ([Formula: see text]exp) was higher than that estimated in phocids, but lower than that reported in cetaceans and the California sea lion. [Formula: see text]exp was maintained over all lung volumes during spontaneous and active respiratory manoeuvres. We conclude that location (water or land) affects lung function in the walrus and should be considered when studying respiratory physiology in semi-aquatic marine mammals.

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.

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 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.