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PubMed2001

Envelope structure of Synechococcus sp. WH8113, a nonflagellated swimming cyanobacterium.

BACKGROUND: Many bacteria swim by rotating helical flagellar filaments. Waterbury et al. discovered an exception, strains of the cyanobacterium Synechococcus that swim without flagella or visible changes in shape. Other species of cyanobacteria glide on surfaces. The hypothesis that Synechococcus might swim using traveling surface waves prompted this investigation. RESULTS: Using quick-freeze electron microscopy, we have identified a crystalline surface layer that encloses the outer membrane of the motile strain Synechococcus sp. WH8113, the components of which are arranged in a rhomboid lattice. Spicules emerge in profusion from the layer and extend up to 150 nm into the surrounding fluid. These spicules also send extensions inwards to the inner cell membrane where motility is powered by an ion-motive force. CONCLUSION: The envelope structure of Synechococcus sp. WH8113 provides new constraints on its motile mechanism. The spicules are well positioned to transduce energy at the cell membrane into mechanical work at the cell surface. One model is that an unidentified motor embedded in the cell membrane utilizes the spicules as oars to generate a traveling wave external to the surface layer in the manner of ciliated eukaryotes.

Cyanobacteria↗
PubMed2004

Swimming gaits, passive drag and buoyancy of diving sperm whales Physeter macrocephalus.

Drag and buoyancy are two primary external forces acting on diving marine mammals. The strength of these forces modulates the energetic cost of movement and may influence swimming style (gait). Here we use a high-resolution digital tag to record depth, 3-D orientation, and sounds heard and produced by 23 deep-diving sperm whales in the Ligurian Sea and Gulf of Mexico. Periods of active thrusting versus gliding were identified through analysis of oscillations measured by a 3-axis accelerometer. Accelerations during 382 ascent glides of five whales (which made two or more steep ascents and for which we obtained a measurement of length) were strongly affected by depth and speed at Reynold's numbers of 1.4-2.8x10(7). The accelerations fit a model of drag, air buoyancy and tissue buoyancy forces with an r(2) of 99.1-99.8% for each whale. The model provided estimates (mean +/- S.D.) of the drag coefficient (0.00306+/-0.00015), air carried from the surface (26.4+/-3.9 l kg(-3) mass), and tissue density (1030+/-0.8 kg m(-3)) of these five animals. The model predicts strong positive buoyancy forces in the top 100 m of the water column, decreasing to near neutral buoyancy at 250-850 m. Mean descent speeds (1.45+/-0.19 m s(-1)) were slower than ascent speeds (1.63+/-0.22 m s(-1)), even though sperm whales stroked steadily (glides 5.3+/-6.3%) throughout descents and employed predominantly stroke-and-glide swimming (glides 37.7+/-16.4%) during ascents. Whales glided more during portions of dives when buoyancy aided their movement, and whales that glided more during ascent glided less during descent (and vice versa), supporting the hypothesis that buoyancy influences behavioural swimming decisions. One whale rested at approximately 10 m depth for more than 10 min without fluking, regulating its buoyancy by releasing air bubbles.

Acceleration↗
PubMed2004

Stroke patterns and regulation of swim speed and energy cost in free-ranging Brünnich's guillemots.

Loggers were attached to free-ranging Brünnich's guillemots Uria lomvia during dives, to measure swim speeds, body angles, stroke rates, stroke and glide durations, and acceleration patterns within strokes, and the data were used to model the mechanical costs of propelling the body fuselage (head and trunk excluding wings). During vertical dives to 102-135 m, guillemots regulated their speed during descent and much of ascent to about 1.6+/-0.2 m s(-1). Stroke rate declined very gradually with depth, with little or no gliding between strokes. Entire strokes from 2 m to 20 m depth had similar forward thrust on upstroke vs downstroke, whereas at deeper depths and during horizontal swimming there was much greater thrust on the downstroke. Despite this distinct transition, these differences had small effect (<6%) on our estimates of mechanical cost to propel the body fuselage, which did not include drag of the wings. Work stroke(-1) was quite high as speed increased dramatically in the first 5 m of descent against high buoyancy. Thereafter, speed and associated drag increased gradually as buoyancy slowly declined, so that mechanical work stroke(-1) during the rest of descent stayed relatively constant. Similar work stroke(-1) was maintained during non-pursuit swimming at the bottom, and during powered ascent to the depth of neutral buoyancy (about 71 m). Even with adjustments in respiratory air volume of +/-60%, modeled work against buoyancy was important mainly in the top 15 m of descent, after which almost all work was against drag. Drag was in fact underestimated, as our values did not include enhancement of drag by altered flow around active swimmers. With increasing buoyancy during ascent above 71 m, stroke rate, glide periods, stroke acceleration patterns, body angle and work stroke(-1) were far more variable than during descent; however, mean speed remained fairly constant until buoyancy increased rapidly near the surface. For dives to depths >20 m, drag is by far the main component of mechanical work for these diving birds, and speed may be regulated to keep work against drag within a relatively narrow range.

Acceleration↗
PubMed2006

Median fin function in bluegill sunfish Lepomis macrochirus: streamwise vortex structure during steady swimming.

Fishes have an enormous diversity of body shapes and fin morphologies. From a hydrodynamic standpoint, the functional significance of this diversity is poorly understood, largely because the three-dimensional flow around swimming fish is almost completely unknown. Fully three-dimensional volumetric flow measurements are not currently feasible, but measurements in multiple transverse planes along the body can illuminate many of the important flow features. In this study, I analyze flow in the transverse plane at a range of positions around bluegill sunfish Lepomis macrochirus, from the trailing edges of the dorsal and anal fins to the near wake. Simultaneous particle image velocimetry and kinematic measurements were performed during swimming at 1.2 body lengths s(-1) to describe the streamwise vortex structure, to quantify the contributions of each fin to the vortex wake, and to assess the importance of three-dimensional flow effects in swimming. Sunfish produce streamwise vortices from at least eight distinct places, including both the dorsal and ventral margins of the soft dorsal and anal fins, and the tips and central notched region of the caudal fin. I propose a three-dimensional structure of the vortex wake in which these vortices from the caudal notch are elongated by the dorso-ventral cupping motion of the tail, producing a structure like a hairpin vortex in the caudal fin vortex ring. Vortices from the dorsal and anal fin persist into the wake, probably linking up with the caudal fin vortices. These dorsal and anal fin vortices do not differ significantly in circulation from the two caudal fin tip vortices. Because the circulations are equal and the length of the trailing edge of the caudal fin is approximately equal to the combined trailing edge length of the dorsal and anal fins, I argue that the two anterior median fins produce a total force that is comparable to that of the caudal fin. To provide additional detail on how different positions contribute to total force along the posterior body, the change in vortex circulation as flow passes down the body is also analyzed. The posterior half of the caudal fin and the dorsal and anal fins add vortex circulation to the flow, but circulation appears to decrease around the peduncle and anterior caudal fin. Kinematic measurements indicate that the tail is angled correctly to enhance thrust through this interaction. Finally, the degree to which the caudal fin acts like a idealized two-dimensional plate is examined: approximately 25% of the flow near the tail is accelerated up and down, rather than laterally, producing wasted momentum, a loss not present in ideal two-dimensional theories.

Animal Structures↗
PubMed2006

Biomimetic evolutionary analysis: testing the adaptive value of vertebrate tail stiffness in autonomous swimming robots.

For early vertebrates, a long-standing hypothesis is that vertebrae evolved as a locomotor adaptation, stiffening the body axis and enhancing swimming performance. While supported by biomechanical data, this hypothesis has not been tested using an evolutionary approach. We did so by extending biomimetic evolutionary analysis (BEA), which builds physical simulations of extinct systems, to include use of autonomous robots as proxies of early vertebrates competing in a forage navigation task. Modeled after free-swimming larvae of sea squirts (Chordata, Urochordata), three robotic tadpoles (;Tadros'), each with a propulsive tail bearing a biomimetic notochord of variable spring stiffness, k (N m(-1)), searched for, oriented to, and orbited in two dimensions around a light source. Within each of ten generations, we selected for increased swimming speed, U (m s(-1)) and decreased time to the light source, t (s), average distance from the source, R (m) and wobble maneuvering, W (rad s(-2)). In software simulation, we coded two quantitative trait loci (QTL) that determine k: bending modulus, E (Nm(-2)) and length, L (m). Both QTL were mutated during replication, independently assorted during meiosis and, as haploid gametes, entered into the gene pool in proportion to parental fitness. After random mating created three new diploid genotypes, we fabricated three new offspring tails. In the presence of both selection and chance events (mutation, genetic drift), the phenotypic means of this small population evolved. The classic hypothesis was supported in that k was positively correlated (r(2)=0.40) with navigational prowess, NP, the dimensionless ratio of U to the product of R, t and W. However, the plausible adaptive scenario, even in this simplified system, is more complex, since the remaining variance in NP was correlated with the residuals of R and U taken with respect to k, suggesting that changes in k alone are insufficient to explain the evolution of NP.

Animals↗
PubMed1983

Locomotion in the North American mink, a semi-aquatic mammal. I. Swimming energetics and body drag.

Oxygen consumption (VO2) during surface swimming and total body drag were investigated in the North American mink, Mustela vision Schreber. Over the range of 0.13-0.70 ms-1, VO2 increased curvilinearly with speed for minks swimming against a current in a water flume. Similarly, body drag of a mink carcass increased non-linearly with speed and was described by the equation, Drag = 1.24 velocity. A streamlined body shape, characteristic of many mustelids, aided in reducing drag at high speeds. Net swimming efficiencies were comparatively low (less than 2.8%) and were attributed to hig levels of drag when on the water surface and the absence of appendage specialization for aquatic locomotion. This lack of specialization probably contributes to high energetic costs but enables the mink to forage in both the aquatic and terrestrial environments.

Animals↗
PubMed1987

Effect of burst swimming and adrenaline infusion on O2 consumption and CO2 excretion in rainbow trout, Salmo gairdneri.

1. Immediately following burst swimming, the oxygen consumption of rainbow trout increased by 71%, carbon dioxide excretion by 104% and the respiratory exchange ratio by 17%. 80 min after burst swimming all of these parameters had returned to levels which were not significantly different from control values. 2. Infusion of adrenaline into resting fish had no significant effect on oxygen consumption or carbon dioxide excretion and therefore there was no significant change in the respiratory exchange ratio. 3. This infusion of adrenaline caused a significant elevation in the red blood cell pH which was still present 80 min later. 4. The present results contrast with those of van den Thillart, Randall & Lin (1983), who demonstrated carbon dioxide retention after burst swimming. While it is possible that catecholamines may inhibit bicarbonate flux through the red blood cell, our experiments indicate that this inhibition would not result in detectable changes in carbon dioxide excretion or, therefore, in the respiratory exchange ratio.

Acclimatization↗
PubMed1988

Regional distribution of blood flow during swimming in the tufted duck (Aythya fuligula).

The distribution of blood flow to a number of organs and tissues of the tufted duck was determined (by the microsphere technique) before and while the birds were swimming at close to their maximum sustainable velocity (i.e. at 0.69 +/- 0.01 ms-1). During swimming, oxygen uptake was twice the pre-exercise value. Cardiac output increased by 70%, there was no significant change in arterial blood pressure and total systemic conductance increased by 44%. There were no significant changes in blood flow to the brain, liver, adrenal glands, spleen and respiratory muscles. Not surprisingly, there were increases in blood flow to the heart (30% increase) and to the muscles of the hindlimbs (to 3.1 times the pre-exercise value). Significant reductions in flow occurred to various parts of the gastrointestinal tract (although not to the gastrointestinal tract as a whole), to the pancreas and to the pectoralis muscles. In the case of the flight musculature as a whole, the reduction was to approximately 40% of the values in the ducks before exercise. Thus, despite the fact that cardiac output was some three times lower than it would have been during flight, there was a clear redistribution of blood away from some visceral organs and inactive muscles during surface swimming in the tufted duck. This lends support to the suggestion that blood is selectively directed to the legs, as well as to the brain and central nervous system (CNS) and away from the visceral organs and inactive muscles during voluntary diving in these birds.

Animals↗
PubMed1991

GTP-gamma-S increases the duration of backward swimming behavior and the calcium action potential in marine Paramecium.

Behavioral and electrophysiological experiments were made to examine the hypothesis that G-proteins modulate the voltage-dependent calcium channel in the marine ciliate Paramecium calkinsi. It was found that guanosine-5'-O-(3-thiotriphosphate) (GTP-gamma-S), an analogue of GTP that binds to and activates G-proteins, increased the duration of backward swimming behavior in reversibly permeabilized Paramecium in an irreversible and concentration-dependent manner. At 1 mumol l-1 GTP-gamma-S, the duration of backward swimming behavior was increased fivefold. Other nucleotides and related compounds did not have a significant effect on the backward swimming behavior. To evaluate whether the behavioral effects were due to ion channel modulation, the calcium action potential in intact Paramecium was monitored before and after guanine nucleotide injection. Within 5 min after the injection of GTP-gamma-S or GTP into the cell, the duration of the calcium action potential was prolonged at least threefold. Like the behavioral response, the GTP-gamma-S effect on the calcium action potential duration was irreversible, whereas the effect of GTP began to decay after 6 min. GDP-beta-S, which binds to and inactivates G-proteins, markedly reduced the calcium action potential within 5 min after injection. These results support the hypothesis that the voltage-dependent calcium channels present in Paramecium are modulated by GTP-binding proteins.

Action Potentials↗
PubMed1991

Effect of endurance swimming on the lactate kinetics of rainbow trout.

The lactate turnover rate of rainbow trout (Oncorhynchus mykiss) was measured by bolus injection of [U-14C]lactate at rest and during prolonged swimming at 85% Ucrit to determine the importance of this metabolic fuel for endurance locomotion in fish, to assess whether lactate exchange between white and red muscle could be a possible mechanism for supplying oxidizable fuel to their lateral red muscle, and to compare the contribution of lactate to total energy provision between teleost and mammalian species. Turnover rate only increased from 4.41 +/- 0.33 to 9.71 +/- 1.69 mumol kg-1 min-1 between rest and prolonged swimming, and the contribution of lactate oxidation to total metabolism declined during exercise. Lactate exchange between white and red muscle is, therefore, not a significant mechanism to fuel the active lateral red musculature during prolonged swimming. The lactate turnover rate of teleosts is one or two orders of magnitude lower than in mammals of equivalent size, but lactate has the same importance as a fuel in both vertebrate groups. However, lactate turnover rate and oxidation rate do not scale with body mass in the same fashion as does metabolic rate. The slope of the mammalian relationship for whole-body lactate turnover and oxidation is much lower (0.58) than the slope of the classic relationship for metabolic rate (0.75), indicating that lactate is a much more important oxidative substrate for small than for large animals.

Animals↗
PubMed1992

Heart rates and swim speeds of emperor penguins diving under sea ice.

Heart rate during overnight rest and while diving were recorded from five emperor penguins with a microprocessor-controlled submersible recorder. Heart rate, cardiac output and stroke volume were also measured in two resting emperor penguins using standard electrocardiography and thermodilution measurements. Swim velocities from eight birds were obtained with the submersible recorder. The resting average of the mean heart rates was 72 beats min-1. Diving heart rates were about 15% lower than resting rates. Cardiac outputs of 1.9-2.9 ml kg-1 s-1 and stroke volumes of 1.6-2.7 ml kg-1 were similar to values recorded from mammals of the same body mass. Swim velocities averaged 3 m s-1. The swim speeds and heart rates suggest that muscle O2 depletion must occur frequently: therefore, many dives require a significant energy contribution from anaerobic glycolysis.

Animals↗
PubMed1993

Neuromuscular organization in the swimming system of the pteropod mollusc Clione limacina.

Swim motor neurons of the pteropod mollusc Clione limacina were identified by a combination of electrophysiological and morphological characteristics. Two types of motor neurons were found, including small motor neurons which are active during both slow and fast swimming and which innervated restricted fields of the ipsilateral wing. General excitor motor neurons have large cell bodies, innervate widespread fields and are recruited into activity for fast swimming. Small motor neurons monosynaptically innervate slow-twitch muscle cells, whereas general excitors monosynaptically innervate both slow-twitch and fast-twitch muscle cells. Activity in general excitors can centrally enhance that in small motor neurons because the neurons are electrically coupled. Neuromuscular recordings and lesion experiments indicate that a peripheral nerve network does not appear to play an important role in the spread of excitation throughout the muscle fields.

Animals↗
PubMed1999

Regulation of swimming in the Culex pipiens (Diptera, Culicidae) pupa: kinematics and locomotory trajectories.

High-speed videography was used to investigate swimming kinematics and locomotory trajectories during escape responses in the pupa of Culex pipiens (Diptera, Culicidae). The pupa can perform straight-line motion despite undergoing backward and forward somersaults through 400 degrees at a rate of 20 s(-)(1). For linear motion to occur, the effective stroke of the abdomen must be delivered during the part of each cycle when the body is facing forwards, allowing passive rotation to carry the body round to the correct position for the start of the next half-stroke. On-off motion of the abdomen is regulated by a click mechanism based on stressed integumentary plates which buckle at strategic points in the cycle. The importance of self-inertia as a component in positional regulation was demonstrated by increasing the kinematic viscosity of the medium: this retarded rotation and resulted in somersaulting on the spot. Whereas normally the pupa invariably directs its swimming downwards, reversal of the usual light gradient produced upward paths as well. It is concluded that straight-line motion is generated automatically by the locomotory machine without any need for sensory feedback about body orientation during each cycle, but that swimming direction can be influenced by sensory cues such as light.

Animals↗
PubMed2001

Swimming speeds and buoyancy compensation of migrating adult chum salmon Oncorhynchus keta revealed by speed/depth/acceleration data logger.

Although the homing migration of Pacific salmon is well documented, the swimming behaviour of the returning salmon has been poorly described, principally as a result of the difficulties encountered in monitoring salmon behaviour in the sea. The present study describes the use of a recently developed electronic data logger to obtain simultaneous recordings of the swimming speed, depth, fin-beating activity and body angle of free-ranging chum salmon Oncorhynchus keta during their homing migration in coastal waters. Chum salmon migrated horizontally at speeds of 1.5-3.0 km h(-1). The gross horizontal distance salmon moved during total recording periods were 1.24- to 19.0-fold greater than the net distance from the release site to the retrieval points. It is suggested that homing salmon did not drift passively but swam actively to the spawning grounds. Salmon preferred the surface water, but also made frequent vertical migrations. The travelled depth of each salmon ranged from 0.36 to 0.64 km per hour. Salmon descended at faster rates and steeper angles than they ascended. Both tailbeat frequency and tail thrust were higher during the ascent than the descent phase. These results suggest that chum salmon spent more energy during the ascent than the descent phase. Profiles of descent rate assumed an arched shape with respect to a change in hydrostatic pressure, while ascent rate increased with decreasing depth. High tailbeat frequencies were found during the course of ascent, which suggests that the salmon did not regulate the volume of air in the swim bladder during short-term vertical migrations.

Animal Migration↗
PubMed2001

Why do cubomedusae have only four swim pacemakers?

The classic view of swimming control in scyphozoan and cubozoan jellyfish involves a diffuse motor nerve net activated by multiple pacemaker sites that interact in a simple resetting hierarchy. Earlier modeling studies of jellyfish swimming, utilizing resetting linkages of multiple pacemakers, indicated that increases in pacemaker number were correlated with increases in the rate and regularity of network activity. We conducted a similar study using the cubozoan jellyfish Carybdea marsupialis, concentrating not only on the adaptive features of multiple pacemaker networks but also on the mechanism of pacemaker interaction. The best fit for our experimental data is a model in which pacemakers express a degree of independence. Thus, our results challenge the idea that pacemaker interactions in scyphozoan and cubozoan medusae are based on a strict resetting hierarchy. Furthermore, our data suggest that the combination of semi-independent linkage of pacemakers with the small pacemaker number characteristic of cubomedusae is important in (i) maintaining a biphasic modulatory capability in the swimming system, and (ii) allowing behaviorally appropriate directional responses to asymmetrical sensory inputs in the radially arranged jellyfish nervous system.

Animals↗
PubMed2002

The notochord of hagfish Myxine glutinosa: visco-elastic properties and mechanical functions during steady swimming.

To determine the possible locomotor functions of the hagfish notochord, we measured its flexural stiffness EI (N m(-2)) and flexural damping C (kg m(3) s(-1)), under in vitro conditions that mimicked the body curvature and bending frequency measured during steady undulatory swimming. To assess the notochord's contribution to the mechanical behavior of the whole body, we also measured EI and C of the whole body, the body with skin removed, and the notochord with the outer fibrous sheath removed. When subjected to dynamic bending at angular frequencies from pi to 6pi rad s(-1) and midline curvatures from 11 to 40 m(-1), 1 cm in situ body segments (N=4), located at an axial position of 37% of the body length, showed significant changes in EI, C, the Young's modulus or material stiffness (E, MPa), the net work to bend the body over a cycle (W, J) and resilience (R, % energy return). When skin, muscles and the outer fibrous sheath of the notochord were removed sequentially, each structural reduction yielded significant changes in mechanical properties: C decreased when the skin was removed, E increased when the muscles were removed, and EI and R decreased when the outer fibrous sheath was removed. Although occupying only a small portion of the cross-sectional area, the notochord provides the body with 75% of its total EI and 80% of total C, by virtue of its high E, ranging from 4 to 8 MPa, which is an order of magnitude greater than that of the whole body. Thus, as the body's primary source of EI and C, the notochord determines the passive (i.e. internal, non-muscular) mechanical behavior of the swimming hagfish. EI and C covary inversely and non-linearly such that as C increases, EI decreases. However, the bending moments M (Nm) produced by each property increase proportionally, and the ratio of stiffness to damping moments, also known as the amplification ratio at resonance, is nearly invariant (approximately 7) with changes in driving frequency. If the body operates in life at or near resonance, the variables EI and C interact over a range of swimming speeds to produce passive mechanical stability.

Animals↗
PubMed1982

Energetic advantages of burst-and-coast swimming of fish at high speeds.

A theoretical model describes how an intermittent swimming style can be energetically advantageous over continuous swimming at high average velocities. Kinematic data are collected from high-speed ciné pictures of free swimming cod and saithe at high velocities in a burst-and-coast style. These data suggest that fish make use of the advantages shown by choosing initial and final burst velocities close to predicted optimal values. The limiting role of rapid glycogen depletion in fast white anaerobic muscle fibres is discussed.

Anaerobiosis↗
PubMed1996

Pharmacological study on Agkistrodon blomhoffii blomhoffii BOIE. V. anti-fatigue effect of the 50% ethanol extract in acute weight-loaded forced swimming-treated rats.

The anti-fatigue effect of 50% ethanol extract ([M]) from the dried whole body of Agkistrodon blomhoffii blomhoffii Boie, was investigated using an acute weight-loaded forced swimming (AWLFS) test by monitoring swimming times, blood biochemical parameters, thiobarbiturate-reactive substances (TBARS) as an index of lipid peroxide and antioxidative enzyme activities in blood and tissue. [M] (500 mg/kg/d), given orally for three successive days, significantly prolonged swimming times. It also inhibited the elevation of TBARS in plasma, liver, brain, kidney and soleus, and inhibited the lowering of catalase activity in erythrocyte, liver and soleus. However, it had no inhibitory effect on the elevation of creatine-kinase activity, free fatty acid and lactic acid levels or on the decrease in glucose level in serum. Also, it decreased the plasma TBARS level and increased the superoxide dismutase activity of plasma and erythrocytes in normal rats. From these results, it can be considered that [M] has an anti-fatigue effect.

Animals↗