Subsurface swimming and stationary diving are metabolically cheap in adult Pacific walruses (Odobenus rosmarus divergens).

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 assessment in the Pacific walrus (Odobenus rosmarus divergens) while resting on land and submerged in water.

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.

Comparative Respiratory Physiology in Cetaceans.

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.

Whistling is metabolically cheap for communicating bottlenose dolphins (Tursiops truncatus).

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% 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 2-min baseline period to establish RMR, followed by a 2-min voluntary resting surface apnea, with or without whistling as cued by the trainers, and then a 5-min 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%) as compared with control trials (-0.5±5.9%; means±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.

Ventilation and gas exchange before and after voluntary static surface breath-holds in clinically healthy bottlenose dolphins, Tursiops truncatus.

We measured respiratory flow (V̇), breathing frequency (fR), tidal volume (VT), breath duration and end-expired O2 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 O2, VT and fR following a breath-hold. The analysis suggests that the dolphins attempt to minimize recovery following a dive by altering VT and fR to rapidly replenish the O2 stores. For the first breath following a surface breath-hold, the end-expired O2 decreased with dive duration, while VT and fR increased. Throughout the recovery period, end-expired O2 increased while the respiratory effort (VT, fR) decreased. We propose that the dolphins alter respiratory effort following a breath-hold according to the reduction in end-expired O2 levels, allowing almost complete recovery after 1.2 min.

Impact of gas emboli and hyperbaric treatment on respiratory function of loggerhead sea turtles (Caretta caretta).

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.