Energetic Cost Of Swimming Research Articles

The contractile efficiency of the mantle muscle of European common cuttlefish (Sepia officinalis) during cyclical contractions.

Escape jet propulsion swimming in cuttlefish (Sepia officinalis) is powered by the circular muscles surrounding the mantle cavity. This mode of locomotion is energetically costly compared with undulatory swimming. The energetic cost of swimming is determined by the mechanical power requirements and the efficiency with which chemical energy is transferred into useful mechanical work. One step in this energy transduction process is the transfer of energy from ATP hydrolysis into mechanical work by the muscles. Here, we determined the efficiency of this step, termed the contractile efficiency. Muscle preparations from the circular muscles of the mantle cavity were subjected to sinusoidal length changes at different cycle frequencies, and stimulated with a phase and duration that maximised initial net work. Changes in ATP, arginine phosphate and octopine content between control and exercised muscles were determined and used to calculate the energy released from ATP hydrolysis (Emet). The maximum contractile efficiency (the ratio of net work to Emet) was 0.37, occurring at the same cycle frequency at which mechanical power was maximal and that was used during jet propulsion swimming, suggesting that cuttlefish muscle is adapted to generate muscular power efficiently. The overall efficiency of cuttlefish jet propulsion swimming was estimated to be 0.17, which is broadly comparable to that measured during animal flight and human-powered pedalled locomotion, indicating the high energetic costs of jet propulsion swimming are not due to inefficient locomotion per se; instead, they result from the relatively high mechanical power requirements.

The effect of particle geometry on squirming through a shear-thinning fluid

Biological and artificial microswimmers often encounter fluid media with non-Newtonian rheological properties. In particular, many biological fluids such as blood and mucus are shear-thinning. Recent studies have demonstrated how shear-thinning rheology can impact substantially the propulsion performance in different manners. In this work, we examine the effect of geometrical shape upon locomotion in a shear-thinning fluid using a prolate spheroidal squirmer model. We use a combination of asymptotic analysis and numerical simulations to quantify how particle geometry impacts the speed and the energetic cost of swimming. The results demonstrate the advantages of spheroidal over spherical swimmers in terms of both swimming speed and energetic efficiency when squirming through a shear-thinning fluid. More generally, the findings suggest the possibility of tuning the swimmer geometry to better exploit non-Newtonian rheological behaviours for more effective locomotion in complex fluids.

Effect of low- and high-carbohydrate diets on swimming economy: a crossover study

BackgroundSwimming economy refers to the rate of energy expenditure relative to swimming speed of movement, is inversely related to the energetic cost of swimming, and is as a key factor influencing endurance swimming performance. The objective of this study was to determine if high-carbohydrate, low-fat (HCLF) and low-carbohydrate, high-fat (LCHF) diets affect energetic cost of submaximal swimming.MethodsEight recreational swimmers consumed two 3-day isoenergetic diets in a crossover design. Diets were tailored to individual food preferences, and macronutrient consumption was 69–16-16% and 16–67-18% carbohydrate-fat-protein for the HCLF and LCHF diets, respectively. Following each 3-day dietary intervention, participants swam in a flume at velocities associated with 50, 60, and 70% of their maximal aerobic capacity (VO2max). Expired breath was collected and analyzed while they swam which enabled calculation of the energetic cost of swimming. A paired t-test compared macronutrient distribution between HCLF and LCHF diets, while repeated-measures ANOVA determined effects of diet and exercise intensity on physiological endpoints.ResultsRespiratory exchange ratio was significantly higher in HCLF compared to LCHF (p = 0.003), but there were no significant differences in the rate of oxygen consumption (p = 0.499) or energetic cost of swimming (p = 0.324) between diets. Heart rate did not differ between diets (p = 0.712), but oxygen pulse, a non-invasive surrogate for stroke volume, was greater following the HCLF diet (p = 0.029).ConclusionsA 3-day high-carbohydrate diet increased carbohydrate utilization but did not affect swimming economy at 50–70% VO2max. As these intensities are applicable to ultramarathon swims, future studies should use higher intensities that would be more relevant to shorter duration events.

Variations in Copepod Proteome and Respiration Rate in Association with Diel Vertical Migration and Circadian Cycle.

The diel vertical migration of zooplankton is a process during which individuals spend the night in surface waters and retreat to depth during the daytime, with substantial implications for carbon transport and the ecology of midwater ecosystems. The physiological consequences of this daily pattern have, however, been poorly studied beyond investigations of speed and the energetic cost of swimming. Many other processes are likely influenced, such as fuel use, energetic trade-offs, underlying diel (circadian) rhythms, and antioxidant responses. Using a new reference transcriptome, proteomic analyses were applied to compare the physiological state of a migratory copepod, Pleuromamma xiphias, immediately after arriving to the surface at night and six hours later. Oxygen consumption was monitored semi-continuously to explore underlying cyclical patterns in metabolic rate under dark-dark conditions. The proteomic analysis suggests a distinct shift in physiology that reflects migratory exertion and changes in metabolism. These proteomic analyses are supported by the respiration experiments, which show an underlying cycle in metabolic rate, with a peak at dawn. This project generates molecular tools (transcriptome and proteome) that will allow for more detailed understanding of the underlying physiological processes that influence and are influenced by diel vertical migration. Further, these studies suggest that P. xiphias is a tractable model for continuing investigations of circadian and diel vertical migration influences on plankton physiology. Previous studies did not account for this cyclic pattern of respiration and may therefore have unrepresented respiratory carbon fluxes from copepods by about 24%.

Viscous propulsion in active transversely isotropic media

Taylor’s swimming sheet is a classical model of microscale propulsion and pumping. Many biological fluids and substances are fibrous, having a preferred direction in their microstructure; for example, cervical mucus is formed of polymer molecules which create an oriented fibrous network. Moreover, suspensions of elongated motile cells produce a form of active oriented matter. To understand how these effects modify viscous propulsion, we extend Taylor’s classical model of small-amplitude zero-Reynolds-number propulsion of a ‘swimming sheet’ via the transversely isotropic fluid model of Ericksen, which is linear in strain rate and possesses a distinguished direction. The energetic costs of swimming are significantly altered by all rheological parameters and the initial fibre angle. Propulsion in a passive transversely isotropic fluid produces an enhanced mean rate of working, independent of the initial fibre orientation, with an approximately linear dependence of the energetic cost on the extensional and shear enhancements to the viscosity caused by fibres. In this regime, the mean swimming velocity is unchanged from the Newtonian case. The effect of the constant term in Ericksen’s model for the stress, which can be identified as a fibre tension or alternatively a stresslet characterising an active fluid, is also considered. This stress introduces an angular dependence and dramatically changes the streamlines and flow field; fibres aligned with the swimming direction increase the energetic demands of the sheet. The constant fibre stress may result in a reversal of the mean swimming velocity and a negative mean rate of working if it is sufficiently large relative to the other rheological parameters.

Mathematical analysis and validation of an exactly solvable model for upstream migration of fish schools in one-dimensional rivers

Upstream migration of fish schools in 1-D rivers as an optimal control problem is formulated where their swimming velocity and the horizontal oblateness are taken as control variables. The objective function to be maximized through a migration process consists of the biological and ecological profit to be gained at the upstream-end of a river, energetic cost of swimming against the flow, and conceptual cost of forming a school. Under simplified conditions where the flow is uniform in both space and time and the profit to be gained at the goal of migration is sufficiently large, the optimal control variables are determined from a system of algebraic equations that can be solved in a cascading manner. Mathematical analysis of the system reveals that the optimal controls are uniquely found and the model is exactly solvable under certain conditions on the functions and parameters, which turn out to be realistic and actually satisfied in experimental fish migration. Identification results of the functional shapes of the functions and the parameters with experimentally observed data of swimming schools of Plecoglossus altivelis (Ayu) validate the present mathematical model from both qualitative and quantitative viewpoints. The present model thus turns out to be consistent with the reality, showing its potential applicability to assessing fish migration in applications.

Adaptation and acclimation of traits associated with swimming capacity in Lake Whitefish (coregonus clupeaformis) ecotypes.

BackgroundImproved performance in a given ecological niche can occur through local adaptation, phenotypic plasticity, or a combination of these mechanisms. Evaluating the relative importance of these two mechanisms is needed to better understand the cause of intra specific polymorphism. In this study, we reared populations of Lake Whitefish (Coregonus clupeaformis) representing the’normal’ (benthic form) and the ‘dwarf’ (derived limnetic form) ecotypes in two different conditions (control and swim-training) to test the relative importance of adaptation and acclimation in the differentiation of traits related to swimming capacity. The dwarf whitefish is a more active swimmer than the normal ecotype, and also has a higher capacity for aerobic energy production in the swimming musculature. We hypothesized that dwarf fish would show changes in morphological and physiological traits consistent with reductions in the energetic costs of swimming and maintenance metabolism.ResultsWe found differences in traits predicted to decrease the costs of prolonged swimming and standard metabolic rate and allow for a more active lifestyle in dwarf whitefish. Dwarf whitefish evolved a more streamlined body shape, predicted to lead to a decreased drag, and a smaller brain, which may decrease their standard metabolic rate. Contrary to predictions, we also found evidence of acclimation in liver size and metabolic enzyme activities.ConclusionResults support the view that local adaptation has contributed to the genetically-based divergence of traits associated with swimming activity. Presence of post-zygotic barriers limiting gene flow between these ecotype pairs may have favoured repeated local adaptation to the limnetic niches.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0732-y) contains supplementary material, which is available to authorized users.

A stochastic bioenergetics model-based approach to translating large river flow and temperature into fish population responses: the pallid sturgeon example

Abstract In managing fish populations, especially at-risk species, realistic mathematical models are needed to help predict population response to potential management actions in the context of environmental conditions and changing climate while effectively incorporating the stochastic nature of real world conditions. We provide a key component of such a model for the endangered pallid sturgeon ( Scaphirhynchus albus ) in the form of an individual-based bioenergetics model influenced not only by temperature but also by flow. This component is based on modification of a known individual-based bioenergetics model through incorporation of: the observed ontogenetic shift in pallid sturgeon diet from marcroinvertebrates to fish; the energetic costs of swimming under flowing-water conditions; and stochasticity. We provide an assessment of how differences in environmental conditions could potentially alter pallid sturgeon growth estimates, using observed temperature and velocity from channelized portions of the Lower Missouri River mainstem. We do this using separate relationships between the proportion of maximum consumption and fork length and swimming cost standard error estimates for fish captured above and below the Kansas River in the Lower Missouri River. Critical to our matching observed growth in the field with predicted growth based on observed environmental conditions was a two-step shift in diet from macroinvertebrates to fish.

Effects of resource distribution on the cost of predator avoidance behaviour in tadpoles

Tadpoles reduce activity and increase hiding in the presence of dragonfly larvae. Several studies demonstrate that tadpoles showing this behaviour have slower growth, however, other studies have found no effect or even positive growth in tadpoles exposed to predators. A recent study demonstrated that swimming is an energetically expensive activity for Bufo arabicus tadpoles. Therefore, if food resources are abundant close to refuges, reduced activity may be an advantage and could offset the cost of reduced foraging. I tested this hypothesis by growing B. arabicus tadpoles with food provided either near or away from shelters, in the presence or absence of caged dragonfly larvae. In the presence of dragonfly larvae, tadpoles provided with food close to shelters were significantly larger than those with food further away. Control tadpoles under both food treatments were intermediate in size, although not statistically different from the predator + near food tadpoles. The results indicate that access to resources is the main determinant of growth in B. arabicus tadpoles and that the energetic cost of swimming is a secondary factor.

Swimming, physical activity and health: a historical perspective

Swimming, which is the coordinated and harmonic movement of the human body inside a liquid medium by means of the combined action of the superior and inferior limbs, is a physical activity which is diffused throughout the whole world and it is practiced by healthy and non-healthy subjects. Swimming is one of the physical activities with less contraindications and, with limited exceptions, can be suggested to individuals of both sexes and of every age range, including the most advanced. Swimming requires energy both for the floating process and for the anterograde progression, with a different and variable osteo-arthro-muscular involvement according to the different styles. The energetic requirement is about four times that for running, with an overall efficiency inferior to 10%; the energetic cost of swimming in the female subject is approximately two thirds of that in the male subject. The moderate aerobic training typical of swimming is useful for diabetic and hypertensive individuals, for people with painful conditions of rachis, as also for obese and orthopaedic patients. Motor activity inside the water reduces the risk of muscular-tendinous lesions and, without loading the joints in excess, requires the harmonic activation of the whole human musculature. Swimming is an activity requiring multiple abilities, ranging from a sense of equilibrium to that of rhythm, from reaction speed to velocity, from joint mobility to resistance. The structured interest for swimming in the perspective of human health from the beginning of civilization, as described in this contribution, underlines the relevance attributed to this activity in the course of human history.

The hydrodynamics of swimming at intermediate Reynolds numbers in the water boatman (Corixidae).

The fluid forces that govern propulsion determine the speed and energetic cost of swimming. These hydrodynamics are scale dependent and it is unclear what forces matter to the tremendous diversity of aquatic animals that are between a millimeter and a centimeter in length. Animals at this scale generally operate within the regime of intermediate Reynolds numbers, where both viscous and inertial fluid forces have the potential to play a role in propulsion. The present study aimed to resolve which forces create thrust and drag in the paddling of the water boatman (Corixidae), an animal that spans much of the intermediate regime (10<Re<200). By measuring the force generated by tethered water boatmen, we found that thrust is generated primarily by drag on the paddling appendages, with a negligible contribution from the acceleration reaction force. Based on these findings, we developed a forward-dynamic model of propulsion in free swimming that accurately predicted changes in the body's center of mass over time. For both tethered and free swimming, we used non-linear optimization algorithms to determine the force coefficients that best matched our measurements. With this approach, the drag coefficients on the body and paddle were found to be up to three times greater than on static structures in fully developed flow at the same Reynolds numbers. This is likely a partial consequence of unsteady interactions between the paddles or between the paddles and the body. In addition, the maximum values for these coefficients were inversely related to the Reynolds number, which suggests that viscous forces additionally play an important role in the hydrodynamics of small water boatmen. This understanding for the major forces that operate at intermediate Reynolds numbers offers a basis for interpreting the mechanics, energetics and functional morphology of swimming in many small aquatic animals.

The rising cost of warming waters: effects of temperature on the cost of swimming in fishes.

Understanding the effects of water temperature on the swimming performance of fishes is central in understanding how fish species will respond to global climate change. Metabolic cost of transport (COT)-a measure of the energy required to swim a given distance-is a key performance parameter linked to many aspects of fish life history. We develop a quantitative model to predict the effect of water temperature on COT. The model facilitates comparisons among species that differ in body size by incorporating the body mass-dependence of COT. Data from 22 fish species support the temperature and mass dependencies of COT predicted by our model, and demonstrate that modest differences in water temperature can result in substantial differences in the energetic cost of swimming.

Differences in the energetic cost of swimming in turbulent flow between wild, farmed and domesticated juvenile Atlantic salmon Salmo salar

Domestication has been shown to have an effect on morphology and behaviour of Atlantic salmon (Salmo salar). We compared swimming costs of three groups of juvenile Atlantic salmon subject to different levels of domestication: (1) wild fish; (2) first generation farmed fish origination from wild genitors; and (2) seventh generation farmed fish originating from Norwegian aquaculture stocks. We assessed swimming costs under two types of turbulent flow (one mean flow velocity of 23 cm s−1 and two standard deviations of flow velocity of 5 and 8 cm s−1). Respirometry experiments were conducted with fish in a mass range of 5–15 g wet at a water temperature of 15° C. Our results confirm (1) that net swimming costs are affected by different levels of turbulence such that, for a given mean flow velocity, fish spent 1·5‐times more energy as turbulence increased, (2) that domesticated fish differed in their morphology (having deeper bodies and smaller fins) and in their net swimming costs (being up to 30·3% higher than for wild fish) and (3) that swimming cost models developed for farmed fish may be also be applied to wild fish in turbulent environments.

On the interaction of turbulent flow and feeding behaviour of juvenile Atlantic Salmon (Salmo salar)

Anthropogenic activities can dramatically modify the riverine habitat of juvenile Atlantic salmon (Salmo salar). In the perspective of protecting and restoring the fluvial habitat, bioenergetic models are often used to estimate fish habitat quality. These models determine the habitat quality as the ratio between the energetic gains (food) and costs (growth, metabolism) of a fish. The energetic costs of swimming in a river are generally estimated using the average flow velocity without consideration of the effect of turbulence. Juvenile Atlantic salmon (JAS) live in rivers characterized by intense velocity fluctuations, which are often described as a succession of high‐ and low‐speed flow regions. These flow structures are likely to affect the JAS activity that consists of long periods of sit‐and‐wait at the top of a protuberant rock interrupted by short bursting motions to capture drifting food particles. To minimize the energetic costs, it is hypothesized that JAS use low‐speed flow regions to initiate and undertake their feeding motions. To improve bioenergetic modelling, this study aimed at analyzing the relation between turbulent flow structures and the feeding behaviour of JAS in a natural gravel‐bed river. We filmed eight JAS during 30 min with a submersible video camera while simultaneously measuring velocity fluctuations close to the fish in the St. Marguerite River, Quebec, Canada. Our results show that the proportion of time used for feeding motions decreases with increasing turbulent intensity and mean flow velocity; and that JAS do not seem to prefer low‐speed flow regions to initiate their feeding motions.

The influence of thermal acclimation on power production during swimming. II. Mechanics of scup red muscle under in vivo conditions.

We have previously shown that the power output of red muscle from warm-acclimated scup is greatly reduced when the fish swim at low temperatures. This reduction occurs primarily because, despite the slowing of muscle relaxation rate at cold temperatures, warm-acclimated scup swim with the same tail-beat frequency and the same stimulation durations, thereby not affording the slower-relaxing muscle any extra time to relax. We hypothesize that power output during swimming could be increased if the stimulus duration were reduced or if the relaxation rate of the red muscle were increased during cold acclimation. Scup were acclimated to 10 degrees C (cold-acclimated) and 20 degrees C (warm-acclimated) for at least 6 weeks. Cold acclimation dramatically increased the ability of scup red muscle to produce power at 10 degrees C. Power output measured from cold-acclimated muscle bundles driven through in vivo conditions measured from cold-acclimated scup swimming at 10 degrees C (i.e. work loops) was generally much greater than that from warm-acclimated muscle driven through its respective in vivo conditions at 10 degrees C. The magnitude of the increase depended both on the anatomical location of the muscle and on swimming speed. Integrated over the length of the fish, the red musculature from cold-acclimated fish generated 2.7, 8.9 and 5.8 times more power than the red musculature from warm-acclimated fish while swimming at 30 cm s(-)(1), 40 cm s(-)(1) and 50 cm s(-)(1), respectively. Our analysis suggests that the cold-acclimated fish should be able to swim in excess of 40 cm s(-)(1) with just their red muscle whereas the warm-acclimated fish must recruit their pink muscle well below this speed. Because the red muscle is more aerobic than the pink muscle, cold acclimation may increase the sustained swimming speed at which scup perform their long seasonal migrations at cool temperatures. We then explored the underlying mechanisms for the increase in muscle power output in cold-acclimated fish. Contrary to our expectations, cold-acclimated muscle did not have a faster relaxation rate; instead, it had an approximately 50 % faster activation rate. Our work-loop studies showed that this faster activation rate, alone, can increase the mechanical power production during cyclical contractions to a surprising extent. By driving cold-acclimated muscle through warm- and cold-acclimated in vivo conditions, we were able to partition the improvement in power production associated with increased activation rate and the approximately 20 % reduction in the duration of electromyographic activity found in the accompanying study. Depending on the position and swimming speed, approximately 60 % of the increase in power output was due to the change in the red muscle's contractile properties (i.e. faster activation); the remainder was due to the shorter stimulus duty cycle of cold-acclimated scup. Thus, by both shortening the in vivo stimulation duration and speeding up the rate of muscle activation as part of cold-acclimation, scup achieve a very large increase in the power output of their red muscle during swimming at low temperature. This increase in power output probably results in an increase in muscle efficiency and, hence, a reduction in the energetic cost of swimming. This increase in power output also reduces reliance on the less aerobic and less fatigue-resistant pink muscle. Both these abilities may increase the swimming speed at which prolonged aerobic muscle activity can occur and thus reduce the travel time for the long seasonal migrations in which scup engage.

Energetic costs of surface swimming and diving of birds.

The energetic costs of swimming at the surface (swimming) and swimming underwater (diving) are compared in tufted ducks (Aythya fuligula) and three species of penguins, the gentoo (Pygoscelis papua), the king (Aptenodytes patagonicus), and the emperor (Aythya forsteri). Ducks swim on the surface and use their webbed feet as paddles, whereas penguins tend to swim just below the surface and use their flippers as hydrofoils, the latter being much more efficient. Penguins are more streamlined in shape. Thus, the amount of energy required to transport a given mass of bird a given distance (known as the cost of transport) is some two to three times greater in ducks than in penguins. Ducks are also very buoyant, and overcoming the force of buoyancy accounts for 60% and 85% of the cost of descent and remaining on the bottom, respectively, in these birds. The energy cost of a tufted duck diving to about 1.7 m is similar to that when it is swimming at its maximum sustainable speed at the surface (i.e., approximately 3.5 times the value when resting on water). Nonetheless, because of the relatively short duration of its dives, the tufted duck dives well within its calculated aerobic dive limit (cADL, usable O(2) stores per rate of O(2) usage when underwater). However, these three species of penguins have maximum dive durations ranging from 5 min to almost 16 min and maximum dive depths from 155 to 530 m. When these birds dive, they have to metabolise at no more than when resting in water in order for cADL to encompass the duration of most of their natural dives. In gentoo and king penguins, there is a fall in abdominal temperature during bouts of diving; this may reduce the oxygen requirements in the abdominal region, thus enabling dive duration to be extended further than would otherwise be the case.

Diving Energetics in King Penguins (Aptenodytes Patagonicus)

Dive duration in wild king penguins and the energetic cost of swimming in a 30m long swim channel were determined at Ile de la Possession, Crozet Archipelago, using external data loggers and respirometry, respectively. Calibrated electronic data loggers equipped with a pressure sensor were used to determine dive durations: 95% of dives were less than 6 min long and 66% of dives were less than 4 min long. Dive patterns show that king penguins may intersperse long dive durations (4-6.3 min) with short ones (1.5-3 min) and make surface pauses of variable duration between them (0.5-3.5 min), or dive regularly (for up to 5 h) with long dive durations (5 min) and constant interdive surface intervals (1.5 min). The latter indicates that the aerobic dive limits (ADL) of this species could be higher and oxygen consumption lower than previously reported. Assuming that king penguins dive within their aerobic limit, different approaches to the analysis of the data obtained in the swim channel are discussed to derive the ADL. Swimming speeds observed in the channel ranged from 0.9 to 3.4 m s-1. Transport costs were lowest between 1.8 and 2.2 m s-1. Although at 2.2 m s-1 king penguins used only 10.3 Wkg-1 over a dive+surface cycle (minimal transport costs of 4.7 J kg-1 m-1), we speculate that tisse oxygen consumption during submergence may be as low as 0.23 ml O2 kg-1 s-1 (2.1 times standard metabolic rate, SMR) or perhaps lower (which gives an ADL of 4.2 min). During surface phases, oxygen uptake would be increased to at least 1 ml O2kg-1 s-1 (9.3 times SMR). This implies that at least 70% of all dives are aerobic. Potential physiological mechanisms allowing king penguins to partition O2 consumption between submergence and surface periods remain, however, unclear.

Travel at low energetic cost by swimming and wave-riding bottlenose dolphins.

Over the past 50 years there has been much speculation about the energetic cost of swimming and wave-riding by dolphins. When aligned properly in front of the bow of moving ships in the stern wake of small boats, on wind waves, and even in the wake of larger cetaceans, the animals appear to move effortlessly through the water without the benefit of propulsive strokes by the flukes. Theoretically, body streamlining as well as other anatomical and behavioural adaptations contribute to low transport costs in these animals. The economy of movement permitted by wave-riding has been perceived as an energetic advantage for the swimming dolphin, but has been hard to prove in the absence of physiological data for exercising cetaceans. Here we determine the aerobic and anaerobic costs of swimming and wave-riding in bottlenose dolphins and find that the minimum cost of transport for swimming dolphins is 1.29 +/- 0.05 J kg-1 m-1 at a cruising speed of 2.1 m s-1. Aerobic costs are nearly twice as high for swimming seals and sea lions, and 8-12 times higher for human swimmers. Wave-riding by dolphins provides additional benefits in terms of speed. The results indicate that behavioural, physiological and morphological factors make swimming an economical form of high-speed travel for dolphins.

Swimming Speed and Oxygen Consumption in the Bathypelagic MysidGnathophausia ingens

The energetic costs of swimming were deter mined for the bathypelagic mysid Gnathophausia in- gens. Individuals over a large size range spontaneously swam at speeds from 5 to 6.5 cm/s. To maintain this speed, smaller animals swam at much higher relative swimming speeds than did larger animals. Routine rates of oxygen consumption were thus considerably higher in the smaller instars. The relationship between standard rates of oxygen consumption and animal size was slightly less than the standard log-log allometric slope of 0.75. Within the speed range of 0-8 cm/s, oxygen consump tion appeared to increase as a linear function of speed. Cost of transport was very high at low speeds. At 5.5 cm/ s, cost of transport was lower than that measured for other crustaceans, but higher than that of fish. Swim ming efficiency increased with speed. While the lower cost of transport and higher swimming efficiency may contribute to G. ingens ' reduced rates of oxygen con sumption as compared to those of shallower-living crus taceans, the major factor appears to be G. ingens' lower

Towards an energy budget for krill: The physiology and biochemistry of Euphausia superba Dana

Published data on the oxygen consumption, nitrogen excretion, feeding, growth and moulting physiology of the Antarctic krill, Euphausia superba, are reviewed with particular reference to experimental methodology and the value of the data for the construction of an energy budget. It is concluded that the relationship between basal oxygen consumption (QO2, in ml h-1) and wet weight (W, in grams) can be described by the relationship QO2=0.0813 W0.88, that filtration rates in krill are high (but possibly reduced in dense swarms), and growth of adult krill in summer is about 2 mm per week. Using these data a preliminary energy budget has been constructed for adult krill at South Georgia in summer. This budget highlights the lack of data on the energetic costs of swimming and feeding in krill, and also the environmental and behavioural data necessary for the extrapolation of an individual energy budget to a swarm or whole population. This preliminary budget suggests a daily energy intake for male krill in summer of about 5% body weight per day. A minimal estimate of the cost of reproduction in female krill from the energy content of the ovary suggests that in a maturing female energy intake can be in excess of 6% body weight per day.