Energetics of swimming of a sea turtle.

Young (mean mass 735 g) green turtles (Chelonia mydas) were able to swim in a water channel at sustained speeds between 0-14 and 0-35 m.s-1. Oxygen consumption at rest was was 0-07 l.kg-1.h-1; at maximum swimming speed oxygen consumption was 3-4 times greater than at rest for a given individual. In comparison with other animals of the same body mass the cost of transport for the green turtle (0.186lO2.kg-1.km-1) is less than that for flying birds but greater than that for fish. From drag measurements it was calculated that the aerobic efficiency of swimming was between 1 and 10%; the higher efficiencies were found at the higher swimming speeds. Based upon the drag calculations for young turtles, it is estimated that adult turtles making the round-trip breeding migration between Brazil and Ascension Island (4800 km) would require the equivalent of about 21% of their body mass in fat stores to account for the energetic cost of swimming.     

Active drag related to velocity in male and female swimmers

Propulsive arm forces of 32 male and 9 female swimmers were measured during front crawl swimming using arms only, in a velocity range between 1.0 m s-1 and 1.8 m s-1. At constant velocity, the measured mean propulsive force Fp equals the mean active drag force (Fd). It was found that Fd is related to the swimming velocity v raised to the power 2.12 +/- 0.20 (males) or 2.28 +/- 0.35 (females). Although many subjects showed rather constant values of Fd/v2, 12 subjects gave significantly (p less than 0.01) stronger or weaker quadratic relationships. Differences in drag force and coefficient of drag between males and females (drag: 28.9 +/- 5.1 N, 20.4 +/- 1.9 N, drag coefficient: 0.64 +/- 0.09, 0.54 +/- 0.07 respectively) are especially apparent at the lowest swimming velocity (1 m s-1), which become less at higher swimming velocities. Possible explanations for the deviation of the power of the velocity from the ideal quadratic dependency are discussed.     

The determination of drag in front crawl swimming

The measurement of drag while swimming (i.e. active drag) is a controversial issue. Therefore, in a group of six elite swimmers two active drag measurement methods were compared to assess whether both measure the same retarding force during swimming. In method 1 push-off forces are measured directly using the system to measure active drag (MAD-system). In method 2 (the velocity perturbation method, VPM) drag is estimated from the difference in swimming speed when subjects swim twice at maximal effort (assuming equal power output and assuming a quadratic drag-speed relationship): once swimming free, and once swimming with a hydrodynamic body attached that created a known additional resistance. The average drag for the VPM tests (53.2 N) was statistically significant and different from the active drag for the MAD-test (66.9 N), paired Student's t-test: 2.484, 12 DF, p=0.029. A post hoc analysis was performed to assess whether the two methods measure a different phenomenon. Based on the drag speed curve obtained with the MAD-system, the VPM-data were re-examined. For diverging drag determinations the assumption of equal power output of the 'free' trial (swimming free) vs. the towing trial (swimming with hydrodynamic buoy) appeared to be violated. The regression of the relative difference in force (MAD vs. VPM) on the relative difference in power (swimming free vs. swimming with hydrodynamic body) was: %Deltadrag=1.898 x %Deltapower -4.498, r2=0.88. This suggests that the major part of the difference in active drag values is due to a non-equal power output in the 'free' relative towing trial during the VPM-test. The simulation of the violation of the equal power output assumption and the calculation of the effect of an other than quadratic drag-speed relationship corroborated the tentative conclusion that both methods measure essentially the same phenomenon and that active drag differences can be explained by a violation of test assumptions.     

The effect of submergence on heart rate and oxygen consumption of swimming seals and sea lions

Respiratory, metabolic, and cardiovascular responses to swimming were examined in two species of pinniped, the harbor seal (Phoca vitulina) and the California sea lion (Zalophus californianus). 1. Harbor seals remained submerged for 82-92% of the time at swimming speeds below 1.2 m.s-1. At higher speeds, including simulated speeds above 1.4 m.s-1, the percentage of time spent submerged decreased, and was inversely related to body weight. In contrast, the percentage of time spent submerged did not change with speed for sea lions swimming from 0.5 m.s-1 to 4.0 m.s-1. 2. During swimming, harbor seals showed a distinct breathhold bradycardia and ventilatory tachycardia that were independent of swimming speed. Average heart rate was 137 beats.min-1 when swimming on the water surface and 50 beats.min-1 when submerged. A bimodal pattern of heart rate also occurred in sea lions, but was not as pronounced as in the seals. 3. The weighted average heart rate (WAHR), calculated from measured heart rate and the percentage time spent on the water surface or submerged, increased linearly with swimming speed for both species. The graded increase in heart rate with exercise load is similar to the response observed for terrestrial mammals. 4. The rate of oxygen consumption increased exponentially with swimming speed in both seals and sea lions. The minimum cost of transport calculated from these rates ranged from 2.3 to 3.6 J.m-1.kg-1, and was 2.5-4.0 times the level predicted for similarly-sized salmonids. Despite different modes of propulsion and physiological responses to swimming, these pinnipeds demonstrate similar transport costs.     

From record performance to hypoxia tolerance: respiratory transition in damselfish larvae settling on a coral reef

The fastest swimming fishes in relation to size are found among coral reef fish larvae on their way to settle on reefs. By testing two damselfishes, Chromis atripectoralis and Pomacentrus amboinensis, we show that the high swimming speeds of the pre-settlement larvae are accompanied by the highest rates of oxygen uptake ever recorded in ectothermic vertebrates. As expected, these high rates of oxygen uptake occur at the cost of poor hypoxia tolerance. However, hypoxia tolerance is needed when coral reef fishes seek nocturnal shelter from predators within coral colonies, which can become severely hypoxic microhabitats at night. When the larvae settle on the reef, we found that they go through a striking respiratory transformation, i.e. the capacity for rapid oxygen uptake falls, while the ability for high-affinity oxygen uptake at low oxygen levels is increased. This transition to hypoxia tolerance is needed when they settle on the reef; this was strengthened by our finding that small resident larvae of Acanthochromis polyacanthus, a damselfish lacking a planktonic larval stage, do not display such a transition, being well adapted to hypoxia and showing relatively low maximum rates of oxygen uptake that change little with age.     

Oxygen uptake during swimming in a hypobaric hypoxic environment

The purpose of this study was to determine oxygen uptake (VO2) at various water flow rates and maximal oxygen uptake (VO2max) during swimming in a hypobaric hypoxic environment. Seven trained swimmers swam in normal [N; 751 mmHg (100.1 kPa)] and hypobaric hypoxic [H; 601 mmHg (80.27 kPa)] environments in a chamber where atmospheric pressure could be regulated. Water flow rate started at 0.80 m.s-1 and was increased by 0.05 m.s-1 every 2 min up to 1.00 m.s-1 and then by 0.05 m.s-1 every minute until exhaustion. At submaximal water flow rates, carbon dioxide production (VCO2), pulmonary ventilation (VE) and tidal volume (VT) were significantly greater in H than in N. There were no significant differences in the response of submaximal VO2, heart rate (fc) or respiratory frequency (fR) between N and H. Maximal VE, fR, VT, fc, blood lactate concentration and water flow rate were not significantly different between N and H. However, VO2max under H [3.65 (SD 0.11) l.min-1] was significantly lower by 12.0% (SD 3.4)% than that in N [4.15 (SD 0.18) l.min-1]. This decrease agrees well with previous investigations that have studied centrally limited exercise, such as running and cycling, under similar levels of hypoxia.     

Energy cost and efficiency of sculling a Venetian gondola

Oxygen uptake was measured on four male subjects during sculling gondolas at constant speeds from approximately 1 to approximately 3 m.s-1. The number of scullers on board in the different trials was one, two or four. Tractional water resistance (drag, D, N) was also measured in the same range of speeds. Energy cost of locomotion per unit of distance (C, J.m-1), as calculated from the ratio of O2 uptake above resting to, increased with v according to a power function (C = 155.2.v1.67; r = 0.88). Also D could be described as a power function of the speed: D = 12.3.v2.21; r = 0.94). The overall efficiency of motion, as obtained from the ratio of D to C, increased with speed from 9.2% at 1.41 m.s-1 to 14.5% at 3.08 m.s-1. It is concluded that, in spite of this relatively low efficiency of motion, the gondola is a very economic means. Indeed, at low speeds (approximately 1 m.s-1), the absolute amount of energy for propelling a gondola is the same as that for waking on the level at the same speed for a subject of 70 kg body mass.     

Respiratory,cardiovascular and metabolic adjustments during steady state swimming in the green turtle,Chelonia mydas

Summary Heart rate and pulmonary artery blood flow of resting green turtles,Chelonia mydas, at 29°C increased with lung ventilation (heart rate from 24±5 to 51±8 beats min). When swimming at 0.6 m s^–1 in water at 30°C, oxygen uptake was 2.83 times and respiratory frequency was 2.75 times the resting values. Heart rate was 1.33 times that during ventilation at rest but 2.83 times that at the end of a breath hold at rest. Partial pressures of oxygen and carbon dioxide, lactic acid concentration and pH of arterial blood, when swimming at 0.5 m s^–1, were similar to those soon after ventilation at rest. Pulmonary blood flow did not decline to low levels between breaths, when the animals were swimming, as it did when they were at rest.In active turtles it appears that pulmonary perfusion remains elevated, supplying oxygen to the locomotory muscles at a sufficiently high rate to support the complete aerobic production of energy, and that respiratory frequency is kept as low as possible, as surfacing for air increases the metabolic cost of swimming.     

Whole-body lift and ground effect during pectoral fin locomotion in the northern spearnose poacher (Agonopsis vulsa)

The northern spearnose poacher, Agonopsis vulsa, is a benthic, heavily armored fish that swims primarily using pectoral fins. High-speed kinematics, whole-body lift measurements, and flow visualization were used to study how A. vulsa overcomes substantial negative buoyancy while generating forward thrust. Kinematics for five freely swimming poachers indicate that individuals tend to swim near the bottom (within 1 cm) with a consistently small (less than 1°) pitch angle of the body. When the poachers swam more than 1 cm above the bottom, however, body pitch angles were higher and varied inversely with speed, suggesting that lift may help overcome negative buoyancy. To determine the contribution of the body to total lift, fins were removed from euthanized fish (n=3) and the lift and drag from the body were measured in a flume. Lift and drag were found to increase with increasing flow velocity and angle of attack (ANCOVA, p<0.0001 for both effects). Lift force from the body was found to supply approximately half of the force necessary to overcome negative buoyancy when the fish were swimming more than 1 cm above the bottom. Lastly, flow visualization experiments were performed to examine the mechanism of lift generation for near-bottom swimming. A vortex in the wake of the pectoral fins was observed to interact strongly with the substratum when the animals approached the bottom. These flow patterns suggest that, when swimming within 1 cm of the bottom, poachers may use hydrodynamic ground effect to augment lift, thereby counteracting negative buoyancy.     

Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus)

Balaenid whales perform long breath-hold foraging dives despite a high drag from their ram filtration of zooplankton. To maximize the volume of prey acquired in a dive with limited oxygen supplies, balaenids must either filter feed only occasionally when prey density is particularly high, or they must swim at slow speeds while filtering to reduce drag and oxygen consumption. Using digital tags with three-axis accelerometers, we studied bowhead whales feeding off West Greenland and present here, to our knowledge, the first detailed data on the kinematics and swimming behaviour of a balaenid whale filter feeding at depth. Bowhead whales employ a continuous fluking gait throughout the bottom phase of foraging dives, moving at very slow speeds (less than 1 m s⁻¹), allowing them to filter feed continuously at depth. Despite the slow speeds, the large mouth aperture provides a water filtration rate of approximately 3 m³ s⁻¹, amounting to some 2000 tonnes of water and prey filtered per dive. We conclude that a food niche of dense, slow-moving zooplankton prey has led balaenids to evolve locomotor and filtering systems adapted to work against a high drag at swimming speeds of less than 0.07 body length s⁻¹ using a continuous fluking gait very different from that of nekton-feeding, aquatic predators.     

Acute cold exposure and the metabolism of glucose and some of its precursors in the liver of the fed and fasted sheep.

Measurements of total body oxygen consumption, visceral and hepatic blood flow, oxygen consumption, exchanges of amino acids, lactate, pyruvate and glucose were made on sheep fed 3--6 h or 21 h before the experiment and exposed for 3 h to a neutral environment (15 degrees C) or a cold environment (0.5 to 4 degrees C with clipped coat and wind speed 2 m.s-1). Recent feeding significantly increasedd the total oxygen consumption and the oxygen consumption of the viscera and liver. No general release of amino acids from the viscera or uptake by the liver after feeding was detected although the arterial plasma concentration of essential amino acids did increase significantly after feeding. The plasma concentration of most non-essential amino acids also increased except that of glycine, which decreased significantly. Cold exposure increased the total oxygen consumption and reduced the respiratory quotient significantly. Release of amino acids from the viscera was stimulated by cold exposure. There was a variable increase in the hepatic uptake of lactate and alanine when the sheep were fasted and cold-exposed. The liver's glucose output doubled and the blood (arterial) glucose concentration significantly increased in the cold.     

Reassessment of the interruption technique for measuring flow resistance in humans

We have previously produced evidence that, in patients with obstructive lung disease, compliance of extrathoracic airways is responsible for lack of mouth-to-alveolar pressure equilibration during respiratory efforts against a closed airway. The flow interruption method for measuring respiratory resistance (Rint) is potentially faced with the same problems. We reassessed the merits of the interruption technique by rendering the extrathoracic airways more rigid and by using a rapid shutter. We measured airway resistance (Raw) with whole body plethysmography during panting (at 2 Hz) and Rint during quiet breathing. Rint and Raw were expressed as specific airway (sGaw) and interruptive conductance (sGint), respectively. In nine healthy subjects (cheeks supported), sGint (0.140 +/- 0.050 s-1.cmH2O-1) was lower (P less than 0.02) than sGaw (0.182 +/- 0.043 s-1.cmH2O-1). By contrast, in 12 patients with severe obstructive lung disease (forced expiratory volume in 1 s/vital capacity = 41.0 +/- 19.8%), sGint (0.058 +/- 0.012 s-1.cmH2O-1) was higher (P less than 0.05) than sGaw (0.047 +/- 0.007 s-1.cmH2O-1), when the cheeks were supported. When the mouth floor was also supported, average values of sGaw (0.048 +/- 0.008 s-1.cmH2O-1) and sGint (0.049 +/- 0.014 s-1.cmH2O-1) became similar. In conclusion, we confirm previous findings in healthy subjects of higher values of Rint, with respect to Raw, probably because of differences in glottis opening between quiet breathing and panting. In airflow obstruction, supporting both the cheeks and the mouth floor decreased sGint, which became similar to sGaw.     

Physiological effects of inspiratory resistance on progressive aerobic work

The purpose of this study was to determine the potential effects on progressive aerobic work while breathing through a new military type chemical and biological (CB) respirator loaded with three different types of purifying canisters. Twelve healthy well-motivated male subjects (mean age 23 +/- 3 years) participated in the study. Results indicated that mean maximal oxygen uptake (VO2max), time to exhaustion, respiratory exchange ratio, rate of perceived exertion, respiratory rate and tidal volume at exhaustion, maximal lactate and the 2-min post-exercise lactate were not significantly influenced when breathing with the respirator and the canisters in comparison to a laboratory valve. Mean pulmonary ventilation, however, was reduced by 21% while oxygen and carbon dioxide ventilatory equivalents were significantly lower by 9% and 8% respectively. Review of the stage-by-stage responses to the treadmill test between the laboratory valve and respirator/canister conditions indicated no significant differences (NS) in oxygen uptake but slightly lower heart rates (NS). Ventilation was not influenced by the canisters until 80% of VO2max at which time the mean oxygen ventilatory equivalent became significantly lower. Blood lactate was significantly depressed between 60% and 90% VO2max under the respirator/canister conditions. It was concluded that, although physiological adaptation occurred, breathing with the new CB respirator and each of the three purifying canisters had no detrimental effect on progressive aerobic work to exhaustion. However, prolonged work at intensities greater than 80-85% of VO2max would in all probability be impaired when breathing with the CB mask and the canisters.     

The aerobic demand of backstroke swimming, and its relation to body size, stroke technique, and performance

Few studies have examined the aerobic demand of backstroke swimming, and its relation to body morphology, technique, or performance. The aims of this study were thus to: i) describe the aerobic demand of backstroke swimming in proficient swimmers at high velocities; ii) assess the effects of body size and stroke technique on submaximal and maximal O2 costs, and; iii) test for a relationship between submaximal O2 costs and maximal performance. Sixteen male competitive swimmers were tested during backstroke swimming at velocities from 1.0 to 1.4 m.s-1. Results showed that VO2 increased linearly with velocity (m.s-1) following the equation VO2 = 6.28v - 3.81 (r = 0.77, SEE/Y = 14.9%). VO2 was also related to the subjects' body mass, height, and armspan. Longer distances per stroke were associated with lower O2 costs, and better maximal performances. A significant relation was found between VO2 at 1.1 m.s-1, adjusted for body mass, and 400 m performance (r = -0.78). Submaximal VO2 was also related to reported times for 100 m and 200 m races. Multiple correlation analyses indicated that VO2 at 1.1 m.s-1 and VO2max accounted for up to 78% of the variance in maximal performances. These results suggest that the assessment of submaximal and maximal VO2 during backstroke swimming may be of value in the training and testing programs of competitive swimmers.     

Methods for the study of amphipod swimming: behavior,morphology, and fluid dynamics

A thorough hydrodynamic approach to the study of swimming in amphipods demands a multipronged attack. A possible first step would be to gather swimming behavior data and determine the biomechanics and kinematics of pleopod beat. This requires careful observation of the swimming modes, swimming speeds, body positions and other aspects of behavior and limb motion that are crucial to swimming. Secondly, it is important to describe the morphology of the body and swimming appendages. Detailed drawings of body shape and design, skeletomusculature, condylic structure, and setal density and distribution on the pleopods and pereopods, are the tools required to ascribe hydrodynamic function to specific limb and body morphology. Finally, the information gathered from behavioral observations bolstered by functional morphology studies is applied to fluid dynamic calculations of drag, lift, and thrust. The theoretical calculations are then compared with empirical determinations of drag, wake generation, vortex shedding frequency, and flow patterns around an amphipod. The fluid dynamic facet of this research is the most challenging and requires an excellent grasp of the fundamental concepts of fluid flow and access to some highly technical equipment. The proposed tripartite approach for the study of amphipod swimming is by no means an exhaustive review of all the techniques that can be employed to quantify amphipod swimming. It will nevertheless permit a rigorous and systematic study of amphipod swimming.     

Rhythms of Locomotion and Oxygen Consumption in the Estuarine Amphipod Corophium Volutator (Crustacea: Amphipoda)

The estuarine amphipod Corophium volutator exhibits an endogenous circatidal rhythm of swimming activity, with maxima occurring just after the expected time of high water, under constant laboratory conditions. Oxygen uptake by Corophium is also subject to modulation across the tidal cycle. The period of highest oxygen uptake occurs during the ebb tide, in phase with the period of maximum swimming activity. A second increase in oxygen uptake during the early flood tide is thought to reflect either in-burrow activity or a previously described rhythm of emergence. This being so, this aspect of the animal's respiratory metabolism may be regulated by an autonomous oscillator independent of that governing the animal's swimming behaviour.     

Is tilting behaviour at low swimming speeds unique to negatively buoyant fish? Observations on steelhead trout,Oncorhynchus mykiss,and bluegill,Lepomis macrochirus

When swimming at low speeds, steelhead trout and bluegill sunfish tilted the body at an angle to the mean swimming direction. Trout swam using continuous body/caudal fin undulation, with a positive (head-up) tilt angle ( 0 , degrees) that decreased with swimming speed ( u , cm s⁻¹) according to: 0 =(164±96).u^{(−1.14±0.41)} (regression coefficients; mean±2 s.e. ). Bluegill swimming gaits were more diverse and negative (head down) tilt angles were usual. Tilt angle was −3·0 ± 0.9° in pectoral fin swimming at speeds of approximately 0.2–1.7 body length s⁻¹ (Ls⁻¹; 3–24 cm s⁻¹), −4.5 ±2.6° during pectoral fin plus body/caudal fin swimming at 1·2–1·7 L s⁻¹ (17–24cm s⁻¹), and −5.0± 1.0° during continuous body/caudal fin swimming at 1.6 and 2.5 L s⁻¹ (22 and 35cm s⁻¹). At higher speeds, bluegill used burst-and-coast swimming for which the tilt angle was 0.1±0.6°. These observations suggest that tilting is a general phenomenon of low speed swimming at which stabilizers lose their effectiveness. Tilting is interpreted as an active compensatory mechanism associated with increased drag and concomitant increased propulsor velocities to provide better stabilizing forces. Increased drag associated with trimming also explains the well-known observation that the relationship between tail-beat frequency and swimming speed does not pass through the origin. Energy dissipated because of the drag increases at low swimming speeds is presumably smaller than that which would occur with unstable swimming.     

Fuel homeostasis in the harbor seal during submerged swimming

1. The turnover rates and oxidation rates of plasma glucose, lactate, and free fatty acids (FFA) were measured in three harbor seals (average mass = 40 kg) at rest or during voluntary submerged swimming in a water flume at 35% (1.3 m.s-1) and 50% (2 m.s-1) of maximum oxygen consumption (MO2max). 2. For seals resting in water, the total turnover rates for glucose, lactate, and FFA were 23.2, 26.2, and 7.5 mumols.min-1.kg-1, respectively. Direct oxidation of these metabolites accounted for approximately 7%, 27%, and 33% of their turnover and 3%, 7%, and 18% of the total ATP production, respectively. 3. For swimming seals, MO2max was achieved at a drag load equivalent to a speed of 3 m.s-1 and averaged 1.85 mmol O2.min-1.kg-1, which is 9-fold greater than resting metabolism in water at 18 degrees C. 4. At 35% and 50% MO2max, glucose turnover and oxidation rates did not change from resting levels. Glucose oxidation contributed about 1% of the total ATP production during swimming. 5. At 50% MO2max, lactate turnover and anaerobic ATP production doubled, but the steady state plasma lactate concentration remained low at 1.1 mM. Lactate oxidation increased 63% but still contributed only 4% of the total ATP production. Anaerobic metabolism contributed about 1% of the total ATP production at rest and during swimming. 6. The plasma FFA concentration and turnover rate increased only 24% and 37% over resting levels, respectively, at 50% MO2max. However, the oxidation rate increased almost 3.5-fold and accounted for 85% of the turnover.(ABSTRACT TRUNCATED AT 250 WORDS)     

Determination and validity of critical velocity as an index of swimming performance in the competitive swimmer.

The purpose of this investigation was to test whether the concept of critical power used in previous studies could be applied to the field of competitive swimming as critical swimming velocity (vcrit). The vcrit, defined as the swimming velocity over a very long period of time without exhaustion, was expressed as the slope of a straight line between swimming distance (dlim) at each speed (with six predetermined speeds) and the duration (tlim). Nine trained college swimmers underwent tests in a swimming flume to measure vcrit at those velocities until the onset of fatigue. A regression analysis of dlim on tlim calculated for each swimmer showed linear relationships (r2 greater than 0.998, P less than 0.01), and the slope coefficient signifying vcrit ranged from 1.062 to 1.262 m.s-1 with a mean of 1.166 (SD 0.052) m.s-1. Maximal oxygen consumption (VO2max), oxygen consumption (VO2) at anaerobic threshold, and the swimming also velocity at the onset of blood lactate accumulation (vOBLA) were also determined during the incremental swimming test. The vcrit showed significant positive correlations with VO2 at anaerobic threshold (r = 0.818, P less than 0.01), vOBLA (r = 0.949, P less than 0.01) and mean velocity of 400 m freestyle (r = 0.864, P less than 0.01). These data suggested that vcrit could be adopted as an index of endurance performance in competitive swimmers.     

Partitioning of oxygen uptake and cost of surfacing during swimming in the air-breathing catfish Pangasianodon hypophthalmus

Though air-breathing has probably evolved mainly as a response to hypoxia, it may provide an important oxygen supplement when metabolism is elevated, as for example during swimming. Due to the increased travelling distance involved when an air-breathing fish swims to and from the surface, and the increased drag when the surface is breached, it can be proposed that air-breathing results in a rise in the apparent cost of transport. In order to investigate this hypothesis, it is necessary to use a fish that is able to swim equally well with and without access to air. The striped catfish Pangasianodon hypophthalmus has been shown to have a sufficiently high capacity for aquatic oxygen uptake in normoxia, to allow for such a comparison. Here, we measured the partitioning of oxygen uptake ( $ \dot{M}{\text{O}}_{2} $ ) during swimming and recovery, and calculated the apparent cost of transport with and without access to air, under normoxic conditions. Aerial $ \dot{M}{\text{O}}_{2} $ constituted 25–40 % of the total $ \dot{M}{\text{O}}_{2} $ during swimming and less than 15 % during recovery. The net cost of transport was 25 % lower in fish that did not air-breathe compared to fish that did, showing that the cost of surfacing can be substantial. This is the first study to measure partitioning in an air-breathing fish during swimming at velocities close to the critical swimming speed.