Swimming suppresses hepatic vitellogenesis in European female silver eels as shown by expression of the estrogen receptor 1, vitellogenin1 and vitellogenin2 in the liver

Background  

When European silver eels (Anguilla anguilla) venture into the Atlantic Ocean for their 6,000 km semelparous spawning run to the Sargasso Sea, they are still in a prepubertal stage. Further sexual development appears to be blocked by dopaminergic inhibition of hypothalamus and pituitary activity. Recently, we found that swimming for several weeks in freshwater stimulated the incorporation of fat droplets in the oocytes. So, it was hypothesized that long term swimming in seawater would release the inhibition further and would also stimulate the production of vitellogenin by the liver.     

Power isn't everything: muscle function and energetic costs during steady swimming in Atlantic cod (Gadus morhua)

Power produced by red myotomal muscles of fish during cruise swimming appears seldom maximized, so we sought to investigate whether economy may impact or dominate muscle function. We measured cost of transport (COT) using oxygen consumption and the strain trajectories and electromyographic activity of red muscle measured at anterior (ANT) and posterior (POST) locations while Atlantic cod (Gadus morhua) swam steadily at speeds between 0.3 and 1.0 body lengths (BL) s(-1). We then measured the power produced by isolated segments of red muscle when activated either as in the swimming cod or such that maximal net power was produced. Patterns of activation during swimming were not optimal for power output and were highly variable between tail beats, particularly at the ANT location and at slow swim speeds. Muscle strain amplitude did not increase until swimming speed reached 0.9 (ANT) versus 0.5 (POST) BL s(-1). These limited power to only 53% (ANT) and 71% (POST) of maximum at slower swim speeds and to 70%-80% of maximum at high swim speeds. COT (resting metabolism subtracted) was minimal at the slowest swim speed, surprisingly, where power was most impaired by activation and strain. Thus, production of powered forces for maneuverability/stability appeared to greatly impact red muscle function during cruise swimming in cod, particularly at slow speeds and in ANT muscle.     

Induced oogenesis of the European eel (Anguilla anguilla L.) in freshwater condition

European eel is a catadromous fish species, which means that after living in freshwater premature individuals adapt to sea water, and migrate to the Sargasso Sea for spawning. Although male eel can be sexually matured even in freshwater, to date, it was believed that female eel can be matured only in seawater. Here we show that the process of sexual maturation may be induced in freshwater by treating female eels with carp pituitary (GSI = 9.87 ± 1.55%). It is thus proposed that seawater condition is not an obligatory environment for stimulating gametogenesis and for artificial maturation of the European eel in neither gender.     

Swimming speed, respiration rate, and estimated cost of transport in adult killer whales

The physiology of free-ranging cetaceans is difficult to study and as a consequence, data on the energetics of these animals are limited. To better understand the energetic cost of swimming in killer whales, total cost of transport ( COT ) was estimated from swimming speeds and respiration rates from wild adult northern resident killer whales ( Orcinus orca ) and reported values of oxygen consumption in captive whales. Respiration rate (breaths per minute) was positively correlated with swimming speed (meters per second), while mass-specific COT (Joules per kilogram per meter) decreased with speed. Lack of data on very fast-swimming animals hindered assessment of the exact speed at which COT was minimal. However, minimum mass-specific COT for killer whales in the present study approached those predicted by a previously published allometric equation for marine mammals, and corresponded to "optimal" swimming speeds of 2.6–3 m/s. Interestingly, the observed average swimming speed (1.6 m/s) was lower than predicted optimal swimming speed. Finally, females with dependent calves had higher respiration rates than females without calves. These findings could be due to synchronous breathing with calves or could result from increased costs of lactation and swimming with a calf in echelon formation. Consequently, females with calves may have much greater COT at optimal swimming speeds than females without calves.     

Swimming performance of silver eels is severely impaired by the swim-bladder parasite Anguillicola crassus

Infection with the swim-bladder parasite Anguillicola crassus is suggested as one of the principal causes of the collapse of the European eel population. This nematode has been introduced in Europe from Asia in the 80s and parasitized in a short time Anguilla eel species in different geographical regions across the globe. The parasites drain energy due to their sanguivorous feeding and they cause mechanical damage on the swim-bladder wall. These two effects are hypothesized to impair the spawning migration of the European eel. In this study, we have investigated both effects on swimming performance. We hypothesized that parasitic sanguivorous activities - related to parasite weight - reduce swimming endurance, while mechanical damage of the swim-bladder impairs buoyancy control. Eighty eels suffering various degrees of infection were introduced in swim-tunnels and subjected to a swimming fitness test. The relation between A. crassus infection and swimming efficiency was measured for large female silver eels swimming at various speeds. Infected eels had lower cruising speeds and a higher cost of transport. Eels without parasites, but with a damaged swim-bladder showed similar effects. Almost half of the eels that contained damaged swim-bladders (43%) stopped swimming at low aerobic swimming speeds (< 0.7 m/s). Simulated migration trials in a recent related study have confirmed that eels with a high parasite level or with damaged swim-bladder show early migration failure (< 1000-km). Reduced swimming performance appears to be associated with swim-bladder dysfunction. As we found that especially silver eels have much higher infection levels than yellow eels, it is concluded that migrating silver eels with severely infected or damaged swim-bladders are unable to reach the spawning grounds.     

Changes in the levels of chloride cells and (Na+ + K+)-dependent ATPase in the gills of yellow and silver eels adapting to seawater.

Changes were measured in the numbers of chloride cells and the levels of (Na+ + K+)-DEPENDENT ATPase in the gills of immature, yellow eels and mature, silver eels during adaptation from freshwater to seawater. The percentage of chloride cells in yellow eels more than doubled after six days in seawater; at this time the specific activity and concentration of (Na+ + K+)-dependent ATPase in gills start to increase in parallel to reach maxima after two weeks that are 2.5 times the starting values. It is concluded that adaptation of yellow eels to seawater involves an increase in the numbers of chloride cells in gills as well as an increased amount of (Na+ + K+)-dependent ATPase per chloride cell. Mature silver eels in freshwater had essentially the same numbers of chloride cells and the same specific activity of the enzyme in the gills as yellow eels fully adapted to seawater. Transferring silver eels to seawater did not alter the percentage of chloride cells in gills although the level of (Na+ + K+)-dependent ATPase and its specific activity increased slightly. Thus, although the silver eel is better prepared for life in seawater than the yellow eel, it still has to attain an increased level of (Na+ + K+)-dependent ATPase in its chloride cells to be fully adapted to seawater.     

Filter-feeding and cruising swimming speeds of basking sharks compared with optimal models: they filter-feed slower than predicted for their size

Movements of six basking sharks (4.0-6.5 m total body length, L(T)) swimming at the surface were tracked and horizontal velocities determined. Sharks were tracked for between 1.8 and 55 min with between 4 and 21 mean speed determinations per shark track. The mean filter-feeding swimming speed was 0.85 m s(-1) (+/-0.05 S.E., n=49 determinations) compared to the non-feeding (cruising) mean speed of 1.08 m s(-1) (+/-0.03 S.E., n=21 determinations). Both absolute (m s(-1)) and specific (L s(-1)) swimming speeds during filter-feeding were significantly lower than when cruise swimming with the mouth closed, indicating basking sharks select speeds approximately 24% lower when engaged in filter-feeding. This reduction in speed during filter-feeding could be a behavioural response to avoid increased drag-induced energy costs associated with feeding at higher speeds. Non-feeding basking sharks (4 m L(T)) cruised at speeds close to, but slightly faster ( approximately 18%) than the optimum speed predicted by the Weihs (1977) [Weihs, D., 1977. Effects of size on the sustained swimming speeds of aquatic organisms. In: Pedley, T.J. (Ed.), Scale Effects in Animal Locomotion. Academic Press, London, pp. 333-338.] optimal cruising speed model. In contrast, filter-feeding basking sharks swam between 29 and 39% slower than the speed predicted by the Weihs and Webb (1983) [Weihs, D., Webb, P.W., 1983. Optimization of locomotion. In: Webb, P.W., Weihs, D. (Eds.), Fish Biomechanics. Praeger, New York, pp. 339-371.] optimal filter-feeding model. This significant under-estimation in observed feeding speed compared to model predictions was most likely accounted for by surface drag effects reducing optimum speeds of tracked sharks, together with inaccurate parameter estimates used in the general model to predict optimal speeds of basking sharks from body size extrapolations.     

Critical swimming speed of yellow‐ and silver‐phase European eel (Anguilla anguilla,L.)

The prime objective of this study was to evaluate differences between the swimming performance of two distinct life stages of European eels. The critical swimming speed (U_crit) of 29 yellow‐ and 33 silver‐phase eels was evaluated in a swim tunnel. Silver‐phase eels showed a better swimming performance (U_crit = 0.66 ms^?1) than yellow individuals (U_crit = 0.43 ms^?1). Male and female silver eels reached an identical U_crit despite their different sizes, which may be a strategy to increase the synchronization of arrival at the spawning grounds.     

Routine and active metabolic rates of migrating adult wild sockeye salmon (Oncorhynchus nerka Walbaum) in seawater and freshwater

We present the first data on the differences in routine and active metabolic rates for sexually maturing migratory adult sockeye salmon (Oncorhynchus nerka) that were intercepted in the ocean and then held in either seawater or freshwater. Routine and active oxygen uptake rates (MO2) were significantly higher (27%-72%) in seawater than in freshwater at all swimming speeds except those approaching critical swimming speed. During a 45-min recovery period, the declining postexercise oxygen uptake remained 58%-73% higher in seawater than in freshwater. When fish performed a second swim test, active metabolic rates again remained 28%-81% higher for fish in seawater except at the critical swimming speed. Despite their differences in metabolic rates, fish in both seawater and freshwater could repeat the swim test and reach a similar maximum oxygen uptake and critical swimming speed as in the first swim test, even without restoring routine metabolic rate between swim tests. Thus, elevated MO2 related to either being in seawater as opposed to freshwater or not being fully recovered from previous exhaustive exercise did not present itself as a metabolic loading that limited either critical swimming performance or maximum MO2. The basis for the difference in metabolic rates of migratory sockeye salmon held in seawater and freshwater is uncertain, but it could include differences in states of nutrition, reproduction, and restlessness, as well as ionic differences. Regardless, this study elucidates some of the metabolic costs involved during the migration of adult salmon from seawater to freshwater, which may have applications for fisheries conservation and management models of energy use.     

Telemetric observations on the spawning migration of the eel (Anguilla anguilla) west of the European continental shelf

Synopsis In the Northern Bay of Biscay and west of the Iberian continental shelf five silver eels (Anguilla anguilla L.) have been tagged with ultrasonic transmitters and tracked 13 to 23 hours over a depth of 200 to 2500 m. Their mean direction from release to the final position of tracking was 288° and significantly closer to the direction of the Sargasso Sea (250° west) than silver eels tracked earlier in the North Sea (341°), possibly 260°. Four of the transmitters were equipped with pressure sensing devices capable of indicating depths of at least 400 m. Three eels tracked at night, during full moon, preferred mean depths of 125, 166 or 215 m. One eel chose a depth of 100 m during moonlight and 50 m after the setting of the moon. Major depth changes, usually occurring one per hour, ranged up to a maximum of 200 m at a maximum vertical speed of 0.6 m sec^–1; this is close to the eels' normal horizontal speed. At dawn all but one dived to a depth of 400 m or more. The eels generally swam below the thermocline and often crossed it.     

Simulating the Oceanic Migration of Silver Japanese Eels

The oceanic migration of silver Japanese eels starts from their continental growth habitats in East Asia and ends at the spawning area near the West Mariana Ridge seamount chain. However, the actual migration routes remain unknown. In this study, we examined the possible oceanic migration routes and strategies of silver Japanese eels using a particle tracking method in which virtual eels (v-eels) were programmed to move vertically and horizontally in an ocean circulation model (Japan Coastal Ocean Predictability Experiment 2, JCOPE2). Four horizontal swimming strategies were tested: random heading, true navigation (readjusted heading), orientation toward the spawning area (fixed heading), and swimming against the Kuroshio. We found that all strategies, except random swimming, allowed v-eels swimming at 0.65 m s⁻¹ to reach the spawning area within eight months after their departure from the south coast of Japan (end of the spawning season). The estimated minimum swimming speed required to reach the area spawning within eight months was 0.1 m s⁻¹ for true navigation, 0.12 m s⁻¹ for constant compass heading, and 0.35 m s⁻¹ for swimming against the Kuroshio. The lowest swimming speed estimated from tracked Japanese eels at sea was 0.03 m.s⁻¹, which would not allow them to reach the spawning area within eight months, through any of the tested orientation strategies. Our numerical experiments also showed that ocean circulation significantly affected the migration of Japanese v-eels. A strong Kuroshio could advect v-eels further eastward. In addition, western Pacific ocean currents accelerated the migration of navigating v-eels. The migration duration was shortened in years with a stronger southward flow, contributed by a stronger recirculation south of Japan, an enhanced subtropical gyre, or a higher southward Kuroshio velocity.     

Two isoforms of the Na+/K+/2Cl- cotransporter are expressed in the European eel (Anguilla anguilla)

Two cDNA isoforms of the NKCC1 secretory cotransporter have been isolated from the European eel. The NKCC1a isoform exhibited mRNA expression in a wide range of tissues in a similar fashion to mammals, whereas NKCC1b was expressed primarily in the brain. The effect of freshwater (FW) to seawater (SW) transfer on NKCC1a expression was dependent on the developmental stage. In non-migratory yellow eels, NKCC1a mRNA expression in the gill was transiently up-regulated 4.3-fold after 2 days but also subsequently by 2.5-6-fold 3 weeks after SW transfer. Gill NKCC1a expression was localised mainly in branchial chloride cells of SW acclimated yellow eels. In contrast to yellow eels, NKCC1a mRNA abundance was not significantly different following SW acclimation in silver eel gill. NKCC1a mRNA abundance decreased in the kidney following SW acclimation and this may correlate with lower tubular ion/fluid secretion and urine flow rates in SW teleosts. Kidney NKCC1a mRNA expression in silver eels was also significantly lower than in yellow eels, suggesting some pre-acclimation of mRNA levels. NKCC1a mRNA was expressed at similar low levels in the middle intestine of FW- and SW-acclimated yellow or silver eels, suggesting the presence of an ion secretory mechanism in this gut segment.     

Male silver eels mature by swimming

Background  

If European silver eels are prevented from reproductive migration, they remain in a prepubertal stage by dopaminergic inhibition of pituitary activity. Because this inhibition is likely a requirement for an extended female growth stage, we tested if it is sex-specific by subjecting both sexes to stimulation by GnRHa (Gonadotropin-Releasing Hormone agonist) – injection or 3-months swimming in seawater.     

Taxonomic revision,ecology and endangerment categorization of the Andean catfish Astroblepus ubidiai (Teleostei: Astroblepidae)

The European eel (Anguilla anguilla Linnaeus 1758) is a species typical for waters of Western Europe. Thanks to early expeditions on the Atlantic Ocean by the Danish biologist Johannes Schmidt who found small (<10mm) leptocephali larvae in the Sargasso Sea about 100 years ago, we have now a strong indication where the spawning site for this species is located. The American eel (Anguilla rostrata, LeSueur) also spawns in the Sargasso Sea. The spawning time and location of both species have been supported and refined in recent analyses of the available historical data. Subsequent ichthyoplankton surveys conducted by McCleave (USA) and Tesch (Germany) in the 1980s indicated an increase in the number of leptocephali <10 mm , confirming and refining the Sargasso Sea theory of Johannes Schmidt. Distinctions between the European and American eel are based on morphological characteristics (number of vertebrae) as well as molecular markers (allozymes, mitochondrial DNA and anonymous genomic-DNA. Although recognised as two distinct species, it remains unclear which mechanisms play a role in species separation during larval drift, and what orientation mechanism eels use during migration in the open sea. The current status of knowledge on these issues will be presented. The hypothesis that all European eel migrate to the Sargasso Sea for reproduction and comprise a single randomly mating population, the so called panmixia theory, was until recently broadly accepted. However, based on field observations, morphological parameters and molecular studies there are some indications that Schmidt’s claim of complete homogeneity of the European eel population and a unique spawning location may be an overstatement. Recent molecular work on European eel indicated a genetic mosaic consisting of several isolated groups, leading to a rejection of the panmixia theory. Nevertheless, the latest extensive genetic survey indicated that the geographical component of genetic structure lacked temporal stability, emphasising the need for temporal replication in the study of highly vagile marine species. Induced spawning of hormone treated eels in the aquarium was collective and simultaneous. In this work for the first time group spawning behaviour has ever been observed and recorded in eels. Studies in swim-tunnels indicate that eels can swim four to six times more efficiently than non-anguilliform fish such as trout. After a laboratory swim trial of eels over 5,500 km, the body composition did not change and fat, protein and carbohydrate were used in the same proportion. This study demonstrated for the first time that European eel are physiologically able of reaching the Sargasso Sea without feeding. Based on catches of newly hatched larvae, temperature preference tests and telemetry tracking of mature hormone treated animals, it can be hypothesised that spawning in the Sargasso Sea is collective and simultaneous, while presumably taking place in the upper 200 m of the ocean. Successful satellite tracking of longfin female eels in New Zealand has been performed to monitor migration pathways. Implementation of this new technology is possible in this species because it is three times larger than the European eel. In the future, miniaturisation of tagging technology may allow European eels to be tracked in time by satellite. The most interesting potential contribution of telemetry tracking of silver eels is additional knowledge about migration routes, rates, and depths. In combination with catches of larvae in the Sargasso Sea, it may elucidate the precise spawning locations of different eel species or groups. Only then, we will be able to define sustainable management issues by integrating this novel knowledge into spawners escapement and juvenile fishing quota.     

Glass eel (Anguilla anguilla) acclimation to freshwater and seawater: morphological changes of the digestive tract

Glass eels ( Anguilla anguilla , L. 1758) caught during ascent at the mouth of the River Tiber were kept in aquaria with freshwater and full strength salinity (35 %) for four months.
Morphological features of glass eels at capture and after four months of experimental rearin are described. The structure of the gut, an important osmoregulatory organ, observed in glass eels in seawater experimental rearing suggests that they undergo an irreversible process of adaptation to freshwater, despite the fact of survival in seawater.     

Kinematics and hydrodynamics of linear acceleration in eels, Anguilla rostrata

The kinematics and hydrodynamics of routine linear accelerations were studied in American eels, Anguilla rostrata, using high-speed video and particle image velocimetry. Eels were examined both during steady swimming at speeds from 0.6 to 1.9 body lengths (L) per second and during accelerations from -1.4 to 1.3 L s(-2). Multiple regression of the acceleration and steady swimming speed on the body kinematics suggests that eels primarily change their tail-tip velocity during acceleration. By contrast, the best predictor of steady swimming speed is body wave speed, keeping tail-tip velocity an approximately constant fraction of the swimming velocity. Thus, during steady swimming, Strouhal number does not vary with speed, remaining close to 0.32, but during acceleration, it deviates from the steady value. The kinematic changes during acceleration are indicated hydrodynamically by axial fluid momentum in the wake. During steady swimming, the wake consists of lateral jets of fluid and has minimal net axial momentum, which reflects a balance between thrust and drag. During acceleration, those jets rotate to point downstream, adding axial momentum to the fluid. The amount of added momentum correlates with the acceleration, but is greater than the necessary inertial force by 2.8+/-0.6 times, indicating a substantial acceleration reaction.     

Influence of oceanic factors on Anguilla anguilla (L.) over the twentieth century in coastal habitats of the Skagerrak,southern Norway

The European eel (Anguilla anguilla L.) is distributed in coastal and inland habitats all over Europe, but spawns in the Sargasso Sea and is thus affected by both continental and oceanic factors. Since the 1980s a steady decline has been observed in the recruitment of glass eels to freshwater and in total eel landings. The eel is considered as critically endangered on the International Union for the Conservation of Nature and Natural Resources Red List of species. The Skagerrak beach seine survey from Norway constitutes the longest fishery-independent dataset on yellow/silver eels (starting in 1904). The Skagerrak coastal region receives larvae born in the Sargasso Sea spawning areas that have followed the Gulf Stream/North Atlantic Drift before they penetrate far into the North Sea. The Skagerrak coastal time series is therefore particularly valuable for exploring the impacts of oceanic factors on fluctuations in eel recruitment abundance. Analyses showed that Sargasso Sea surface temperature was negatively correlated with eel abundance, with a lag of 12 years revealing a cyclic and detrimental effect of high temperatures on the newly hatched larvae. The North Atlantic Oscillation index and inflow of North Atlantic water into the North Sea were negatively correlated with eel abundance, with a lag of 11 years. Increased currents towards the North Atlantic during high North Atlantic Oscillation years may send larvae into the subpolar gyre before they are ready to metamorphose and settle, resulting in low recruitment in the northern part of the distribution area for these years. The Skagerrak time series was compared with glass eel recruitment to freshwater in the Netherlands (Den Oever glass eel time series), and similar patterns were found revealing a cycle linked to changes in oceanic factors affecting glass eel recruitment. The recent decline of eels in the Skagerrak also coincided with previously documented shifts in environmental conditions of the North Sea ecosystem.     

Evidence for energy savings from aerial running in the Svalbard rock ptarmigan (Lagopus muta hyperborea)

Svalbard rock ptarmigans were walked and run upon a treadmill and their energy expenditure measured using respirometry. The ptarmigan used three different gaits: a walking gait at slow speeds (less than or equal to 0.75 m s(-1)), grounded running at intermediate speeds (0.75 m s(-1) < U < 1.67 m s(-1)) and aerial running at high speeds (greater than or equal to 1.67 m s(-1)). Changes of gait were associated with reductions in the gross cost of transport (COT; J kg(-1) m(-1)), providing the first evidence for energy savings with gait change in a small crouched-postured vertebrate. In addition, for the first time (excluding humans) a decrease in absolute metabolic energy expenditure (rate of O(2) consumption) in aerial running when compared with grounded running was identified. The COT versus U curve varies between species and the COT was cheaper during aerial running than grounded running, posing the question of why grounded running should be used at all. Existing explanations (e.g. stability during running over rocky terrain) amount to just so stories with no current evidence to support them. It may be that grounded running is just an artefact of treadmill studies. Research investigating the speeds used by animals in the field is sorely needed.     

流速对细鳞裂腹鱼游泳行为及能量消耗影响的研究

通过自制密封的鱼类游泳实验装置, 研究了流速对细鳞裂腹鱼游泳行为和能量消耗的影响。结果显示,细鳞裂腹鱼的摆尾频率随游泳速度的变化有明显的变化规律, 摆尾频率随着流速的增加而显著性的增加,而摆尾幅度有减小趋势, 差异性不显著。结果还表明, (26±1) ℃时, (10.60±0.54) cm 细鳞裂腹鱼的相对临界游泳速度为(11.5±0.5) BL/s, 绝对临界游泳速度为(110.28±2.02) cm/s。测定的相对临界流速较其他的鲤科鱼大,是对生存水流环境(流速0.5—1.5m/s)适应性的表现。这一结果表明鱼类的游泳能力是能够训练的。运动代谢率与相对流速的关系为, AMR = 93.08e(0.307v) + 314.33, R2= 0.994; 单位距离能耗与流速的指数关系为COT =28e (-1.03V) +6.05, R2= 0.998。流速达到8 BL/s 时, 裂腹鱼耗氧率开始下降, 从流速7 BL/s 时, (1245.57±90.97 )mg O2/(kg·h)最大, 下降到(978.78±189.38) mg O2/(kg·h)。1—7 BL/s 流速范围内, 裂腹鱼单位时间内的耗氧率随着游泳速度的增加而增加, 而且随着游泳速度的增加, 单位距离能耗(COT)逐渐减少, 最小能耗在6 倍体长流速, 0.68 m/s 时, 为(6.00±1.57) J/(kg·m), 其能量利用效率最大。       

Optimal shape and motion of undulatory swimming organisms

Undulatory swimming animals exhibit diverse ranges of body shapes and motion patterns and are often considered as having superior locomotory performance. The extent to which morphological traits of swimming animals have evolved owing to primarily locomotion considerations is, however, not clear. To shed some light on that question, we present here the optimal shape and motion of undulatory swimming organisms obtained by optimizing locomotive performance measures within the framework of a combined hydrodynamical, structural and novel muscular model. We develop a muscular model for periodic muscle contraction which provides relevant kinematic and energetic quantities required to describe swimming. Using an evolutionary algorithm, we performed a multi-objective optimization for achieving maximum sustained swimming speed U and minimum cost of transport (COT)--two conflicting locomotive performance measures that have been conjectured as likely to increase fitness for survival. Starting from an initial population of random characteristics, our results show that, for a range of size scales, fish-like body shapes and motion indeed emerge when U and COT are optimized. Inherent boundary-layer-dependent allometric scaling between body mass and kinematic and energetic quantities of the optimal populations is observed. The trade-off between U and COT affects the geometry, kinematics and energetics of swimming organisms. Our results are corroborated by empirical data from swimming animals over nine orders of magnitude in size, supporting the notion that optimizing U and COT could be the driving force of evolution in many species.