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George V Lauder

Publications and source records attributed to George V Lauder.

16 recordsLinked to original sources
PubMed2006

Ontogeny of form and function: locomotor morphology and drag in zebrafish (Danio rerio).

Many fish species transform in body shape during growth, but it remains unclear how this influences the mechanics of locomotion. Therefore, the present study focused on understanding how drag generation during coasting is affected by ontogenetic changes in the morphology of zebrafish (Danio rerio). The shapes of the body and fins were measured from photographs of fish ranging in size from small larvae to mature adults and these morphometrics were compared to drag coefficients calculated from high-speed video recordings of routine swimming. We found that the viscous drag coefficient of larval and juvenile fish increased by more than an order of magnitude during growth and the inertial drag coefficient decreased at a comparable rate in adults. These hydrodynamic changes occurred as zebrafish disproportionately increased the span of their fins and their body changed shape from elongated to streamlined, as reflected by the logistic growth of a newly defined streamlining index, SL. These results suggest that morphological changes incur a performance cost by generating greater drag when larvae and juveniles operate in the viscous regime, but later provide a performance benefit by reducing pressure drag in the inertial regime of the adult stage.

Animals↗
PubMed2005

The mechanical scaling of coasting in zebrafish (Danio rerio).

Many fish species span two or three orders of magnitude in length during the growth from larvae to adults, and this change may have dramatic consequences for locomotor performance. We measured how the performance of coasting changes over the life history of zebrafish (Danio rerio) and examined the scaling of mechanics underlying this change. Adult zebrafish coast disproportionately further and faster and maintain their speed for a longer duration than do larvae and juveniles. Measurements of drag on tethered dead fish suggest that adult fish operate in an inertial regime by coasting at relatively high Reynolds numbers (Re > 1000), and in vivo drag measurements showed adults to operate with a drag coefficient (Cinert approximately 0.024) that was consistent with previously published estimates. However, drag scaled differently at lower Re values than those assumed in previous studies. We found a viscous regime at Re < 300, which corresponds to the routine coasting of larvae and juveniles. Despite these changes in hydrodynamics over growth, a mathematical model of coasting mechanics suggests that the disproportionately longer coasting of adults is caused primarily by their large body mass and high speed at the beginning of coasting. We therefore propose that changes in coasting performance with growth are dictated primarily by the scaling of momentum rather than resulting from hydrodynamic changes. These results provide an opportunity for new interpretations of function in the growth and evolution of fish.

Animals↗
PubMed2005

Locomotor function of the dorsal fin in rainbow trout: kinematic patterns and hydrodynamic forces.

In this study, we examine the kinematics and hydrodynamics of the soft dorsal fin in a representative basal teleost, the rainbow trout (Oncorhynchus mykiss), during steady rectilinear locomotion at 0.5-2.0 body lengths (L) s(-1) and during maneuvering. During steady swimming, dorsal fin height and sweep amplitude decrease with increasing speed. The dorsal fin wake, as viewed within a horizontal plane, consists of paired vortices on each side of the body (0.5 L s(-1)) or nearly linearly arrayed vortex centers above the body (1.0 L s(-1)) with central jet flows directed predominately laterally (lateral:thrust force ratio = 5-6). At 2.0 L s(-1), the dorsal fin is no longer recruited to add momentum to the wake. This pattern of decreasing involvement of the trout dorsal fin in thrust production with increasing speed contrasts with the results of our previous study of the soft dorsal fin of sunfish (Lepomis), which is hydrodynamically inactive at low speed and sheds a propulsive vortex wake at higher speed. Yawing maneuvers by trout involve unilateral production of a single vortex ring by the dorsal fin with a strong jet flow oriented almost directly laterally. During steady swimming, interception by the tail of the dorsal fin's vortical wake and the adipose fin's non-vortical (drag) wake is hypothesized as a mechanism for enhancing tail thrust. This study provides the first experimental evidence that the plesiomorphic soft dorsal fin of ray-finned fishes acts as an ancillary force generator during axial locomotion. We suggest that the distinction often made between median and paired fin (MPF) propulsion and body and caudal fin (BCF) propulsion in fishes obscures the important role of multiple propulsors acting in a coordinated fashion. Using a combination of anterior median fin oscillation and axial undulation, without continuous paired fin excursions, trout employ an ;M-BCF' gait during steady swimming. The primarily lateral orientation of dorsal fin force in trout induces corresponding roll and yaw moments, which must be countered by forces from the caudal, anal and paired fins. Locomotion in trout therefore involves the simultaneous active use of multiple fins, presumably to maintain body stability in the face of environmental perturbations.

Animal Structures↗
PubMed2005

Constraints on starting and stopping: behavior compensates for reduced pectoral fin area during braking of the bluegill sunfish Lepomis macrochirus.

Many natural animal movements involve accelerating from a standstill and then stopping. Obstacles in natural environments often limit the straight-line distance available for movement, and decreased braking ability theoretically can limit speed for short distances. Consequently, braking ability can be important for avoiding collisions with obstacles and exploiting resources effectively in complex environments. A presumed morphological correlate of improved braking performance in fish is increased pectoral fin area, because most fish protract these structures as they decelerate. However, the kinematics and modulation of velocity during starting and stopping are poorly understood for most species of fish as well as most species of animals. Thus, for bluegill sunfish Lepomis macrochirus with complete and partially ablated pectoral fins (35% original fin area), we analyzed high speed video recordings (200 images s-1) of predatory attacks with a start and stop in a short, standardized distance (40 cm). We quantified body displacement, velocity, acceleration, deceleration and several fin angle variables during each feeding. Unexpectedly, several variables including maximum velocity and maximum deceleration (grand means 72 cm s-1 and -512 cm s-2, respectively) did not change significantly with reduced pectoral fin area. The average values of braking movements of the median and caudal fins did increase with decreased pectoral fin area but lacked statistically significant differences. The primary mechanism of attaining similar braking performance with decreased area of the pectoral fins was that they were protracted significantly more (mean difference=42 degrees) and with a significantly faster average velocity of protraction. Thus, pectoral fin area appears unlikely to be the primary constraint on braking performance for this particular task.

Acceleration↗
PubMed2004

Running on water: Three-dimensional force generation by basilisk lizards.

Water provides a unique challenge for legged locomotion because it readily yields to any applied force. Previous studies have shown that static stability during locomotion is possible only when the center of mass remains within a theoretical region of stability. Running across a highly yielding surface could move the center of mass beyond the edges of the region of stability, potentially leading to tripping or falling. Yet basilisk lizards are proficient water runners, regularly dashing across bodies of water to evade predators. We present here direct measurements of time-averaged force produced by juvenile plumed basilisk lizards (Basiliscus plumifrons) while running across water. By using digital particle image velocimetry to visualize fluid flow induced by foot movement, we show that sufficient support force is generated for a lizard to run across water and that novel strategies are also required to run across a highly yielding surface. Juvenile basilisk lizards produce greatest support and propulsive forces during the first half of the step, when the foot moves primarily vertically downwards into the water; they also produce large transverse reaction forces that change from medial (79% body weight) to lateral (37% body weight) throughout the step. These forces may act to dynamically stabilize the lizards during water running. Our results give insight into the mechanics of how basilisk lizards run across water and, on a broader scale, provide a conceptual basis for how locomotor surface properties can challenge established rules for the mechanics of legged locomotion.

Animals↗
PubMed2004

The hydrodynamics of eel swimming: I. Wake structure.

Eels undulate a larger portion of their bodies while swimming than many other fishes, but the hydrodynamic consequences of this swimming mode are poorly understood. In this study, we examine in detail the hydrodynamics of American eels (Anguilla rostrata) swimming steadily at 1.4 L s(-1) and compare them with previous results from other fishes. We performed high-resolution particle image velocimetry (PIV) to quantify the wake structure, measure the swimming efficiency, and force and power output. The wake consists of jets of fluid that point almost directly laterally, separated by an unstable shear layer that rolls up into two or more vortices over time. Previously, the wake of swimming eels was hypothesized to consist of unlinked vortex rings, resulting from a phase offset between vorticity distributed along the body and vorticity shed at the tail. Our high-resolution flow data suggest that the body anterior to the tail tip produces relatively low vorticity, and instead the wake structure results from the instability of the shear layers separating the lateral jets, reflecting pulses of high vorticity shed at the tail tip. We compare the wake structure to large-amplitude elongated body theory and to a previous computational fluid dynamic model and note several discrepancies between the models and the measured values. The wake of steadily swimming eels differs substantially in structure from the wake of previously studied carangiform fishes in that it lacks any significant downstream flow, previously interpreted as signifying thrust. We infer that the lack of downstream flow results from a spatial and temporal balance of momentum removal (drag) and thrust generated along the body, due to the relatively uniform shape of eels. Carangiform swimmers typically have a narrow caudal peduncle, which probably allows them to separate thrust from drag both spatially and temporally. Eels seem to lack this separation, which may explain why they produce a wake with little downstream momentum while carangiform swimmers produce a wake with a clear thrust signature.

Analysis of Variance↗
PubMed2004

Hydrodynamics of surface swimming in leopard frogs (Rana pipiens).

The kinematics of swimming frogs have been studied extensively in the past and, based on these results, hypotheses regarding the hydrodynamics of frog swimming can be generated. To test these hypotheses we used digital particle image velocimetry (DPIV) to quantify the flow structure of the wake produced by the feet during the propulsion phase of the kick of surface swimming frogs (Rana pipiens). These frogs use two different gaits, asynchronous and synchronous kicking, and the magnitude of the thrust produced by the feet differs between asynchronous (34+/-5.4 mN foot(-1)) and synchronous kicking (71+/-13.3 mN foot(-1)), as does maximum swimming speed, with higher swimming speed and forces produced during the synchronous kicks. Previous studies have suggested that an interaction between the feet, resulting in a single posteriorly directed fluid jet, as the feet come together at the end of synchronous kicks, may augment force production. Our results show, however, that each foot produces its own distinct vortex ring, in both asynchronous and synchronous kicking of the feet. There is no evidence of a central jet being produced even during powerful synchronous kicks (maximum thrust calculated was 264 mN foot(-1)). An alternative mechanism of force production could be the lift-based paddling recently suggested for delta-shaped feet of swimming birds. However, the orientation of the vortex rings generated by the feet is almost perpendicular to the swimming direction for both gaits and there is only a slight asynchrony of the shedding of the distal (start) and proximal (stop) vortex rings, which is different from what would be expected by a dominantly lift-based mechanism. Thus, our results do not support lift as a major mechanism contributing to thrust. Instead, our data support the hypothesis that propulsion is based on drag and acceleration reaction forces where the thrust is generated by separated, but attached, vortex rings on the suction side of the feet, resulting in vortices that are shed behind the frogs during both asynchronous and synchronous kicking.

Acceleration↗
PubMed2003

Fish exploiting vortices decrease muscle activity.

Fishes moving through turbulent flows or in formation are regularly exposed to vortices. Although animals living in fluid environments commonly capture energy from vortices, experimental data on the hydrodynamics and neural control of interactions between fish and vortices are lacking. We used quantitative flow visualization and electromyography to show that trout will adopt a novel mode of locomotion to slalom in between experimentally generated vortices by activating only their anterior axial muscles. Reduced muscle activity during vortex exploitation compared with the activity of fishes engaged in undulatory swimming suggests a decrease in the cost of locomotion and provides a mechanism to understand the patterns of fish distributions in schools and riverine environments.

Animals↗
PubMed2003

Quantification of flow during suction feeding in bluegill sunfish.

Nearly all aquatic-feeding vertebrates use some amount of suction to capture prey items. Suction prey capture occurs by accelerating a volume of water into the mouth and taking a prey item along with it. Yet, until recently, we lacked the necessary techniques and analytical tools to quantify the flow regime generated by feeding fish. We used a new approach; Digital Particle Image Velocimetery (DPIV) to measure several attributes of the flow generated by feeding bluegill sunfish. We found that the temporal pattern of flow was notably compressed during prey capture. Flow velocity increased rapidly to its peak within 20 ms of the onset of the strike, and this peak corresponded to the time that the prey entered the mouth during capture. The rapid acceleration and deceleration of water suggests that timing is critical for the predator in positioning itself relative to the prey so that it can be drawn into the mouth along with the water. We also found that the volume of water affected by suction was spatially limited. Only rarely did we measure significant flow beyond 1.75 cm of the mouth aperture (in 20 cm fish), further emphasizing the importance of mechanisms, like locomotion, that place the fish mouth in close proximity to the prey. We found that the highest flows towards the mouth along the fish midline were generated not immediately in front of the open mouth, but approximately 0.5 cm anterior to the mouth opening. Away from the midline the peak in flow was closer to the mouth. We propose that this pattern indicates the presence of a bow wave created by the locomotor efforts of the fish. In this scheme, the bow wave acts antagonistically to the flow of water generated by suction, the net effect being to push the region of peak flow away from the open mouth. The peak was located farther from the mouth opening in strikes accompanied by faster locomotion, suggesting faster fish created larger bow waves.

Journal Article↗
PubMed2003

Function of pectoral fins in rainbow trout: behavioral repertoire and hydrodynamic forces.

Salmonid fishes (trout, salmon and relatives) have served as a model system for study of the mechanics of aquatic animal locomotion, yet little is known about the function of non-axial propulsors in this major taxonomic group. In this study we examine the behavioral and hydromechanical repertoire of the paired pectoral fins of rainbow trout Oncorhynchus mykiss, performing both steady rectilinear swimming and unsteady maneuvering locomotion. A combination of kinematic analysis and quantitative flow visualization (using digital particle image velocimetry) enables identification of the propulsive roles played by pectoral fin motions. During constant-speed swimming (0.5 and 1.0 body length s(-1)), the pectoral fins remain adducted against the body. These fins are actively recruited, however, for a variety of maneuvering behaviors, including station holding in still water (hovering), low-speed (i.e. non-fast-start) turning, and rapid deceleration of the body during braking. Despite having a shallow pectoral-fin base orientation (the plesiomorphic teleost condition), trout are capable of rotating the fin base over 30 degrees during maneuvering, which affords the fin an impressive degree of kinematic versatility. When hovering, the pectoral fins are depressed beneath the body and twisted along their long axes to allow anteroposterior sculling. During turning and braking, the fins undergo spanwise rotation in the opposite direction and exhibit mediolateral and dorsoventral excursions. Water velocity fields and calculated momentum flows in the wake of the pectoral fins reveal that positive thrust is not generated during maneuvering, except during the retraction half-stroke of hovering. Relatively large laterally directed fluid force (mean 2.7 mN) is developed during turning, whose reaction powers yawing rotation of the body (4-41 degrees s(-1)). During deceleration, the wake-force line of action falls below the center of mass of the body, and this result supports a long-standing mechanical model of braking by fishes with ventrally positioned paired fins. Despite its traditional categorization as a propulsor of limited functional importance, the salmoniform pectoral fin exhibits a diverse locomotor repertoire comparable to that of higher teleostean fishes.

Animals↗
PubMed2003

The Kármán gait: novel body kinematics of rainbow trout swimming in a vortex street.

Most fishes commonly experience unsteady flows and hydrodynamic perturbations during their lifetime. In this study, we provide evidence that rainbow trout Oncorhynchus mykiss voluntarily alter their body kinematics when interacting with vortices present in the environment that are not self-generated. To demonstrate this, we measured axial swimming kinematics in response to changes in known hydrodynamic wake characteristics. We compared trout swimming in the Kármán street behind different diameter cylinders (2.5 and 5 cm) at two flow speeds (2.5 and 4.5 L s(-1), where L is total body length) to trout swimming in the free stream and in the cylinder bow wake. Trout swimming behind cylinders adopt a distinctive, previously undescribed pattern of movement in order to hold station, which we term the Kármán gait. During this gait, body amplitudes and curvatures are much larger than those of trout swimming at an equivalent flow velocity in the absence of a cylinder. Tail-beat frequency is not only lower than might be expected for a trout swimming in the reduced flow behind a cylinder, but also matches the vortex shedding frequency of the cylinder. Therefore, in addition to choosing to be in the slower flow velocity offered behind a cylinder (drafting), trout are also altering their body kinematics to synchronize with the shed vortices (tuning), using a mechanism that may not involve propulsive locomotion. This behavior is most distinctive when cylinder diameter is large relative to fish length. While tuning, trout have a longer body wavelength than the prescribed wake wavelength, indicating that only certain regions of the body may need to be oriented in a consistent manner to the oncoming vortices. Our results suggest that fish can capture energy from vortices generated by the environment to maintain station in downstream flow. Interestingly, trout swimming in front of a cylinder display lower tail-beat amplitudes and body wave speeds than trout subjected to any of the other treatments, implying that the bow wake may be the most energetically favorable region for a fish to hold station near a cylinder.

Animals↗
PubMed2002

Forces, fishes, and fluids: hydrodynamic mechanisms of aquatic locomotion.

Understanding how fishes generate external fluid force to swim steadily and maneuver has proven to be difficult because water does not provide a stable platform for force measurement. But new methods in experimental fluid mechanics provide insights into the physiological mechanisms of aquatic force generation and limits to locomotor performance.

Animals↗
PubMed2002

Hydrodynamics of caudal fin locomotion by chub mackerel, Scomber japonicus (Scombridae).

As members of the derived teleost fish clade Scombridae, mackerel exhibit high-performance aquatic locomotion via oscillation of the homocercal forked caudal fin. We present the first quantitative flow visualization of the wake of a scombrid fish, chub mackerel Scomber japonicus (20-26 cm fork length, FL), swimming steadily in a recirculating flow tank at cruising speeds of 1.2 and 2.2FL s(-1). Thrust was calculated from wake measurements made separately in the horizontal (frontal) plane and vertical (parasagittal) planes using digital particle image velocimetry (DPIV) and compared with drag measurements obtained by towing the same specimens of S. japonicus post mortem. Patterns of flow indicated that the wake consisted of a series of linked elliptical vortex rings, each with central jet flow. The length of the minor axis (height) of the vortex rings was approximately equal to caudal fin span; the length of the major ring axis was dependent on swimming speed and was up to twice the magnitude of ring height. Profiles of wake velocity components were similar to theoretical profiles of vortex rings. Lift, thrust and lateral forces were calculated from DPIV measurements. At 1.2FL s(-1), lift forces measured relative to the X axis were low in magnitude (-1+/-1 mN, mean +/- S.D., N=20) but oriented at a mean angle of 6 degrees to the body axis. Reaction forces tend to rotate the fish about its center of mass, tipping the head down. Thus, the homocercal caudal fin of S. japonicus functions asymmetrically in the vertical plane. Pitching moments may be balanced anteriorly via lift generation by the pectoral fins. Thrust estimates for the two smallest fish based on DPIV analysis were not significantly different from drag measurements made by towing those same animals. At a speed of 1.2FL s(-1), thrust magnitude was 11+/-6 mN (mean +/- S.D, N=40). Lateral force magnitudes were approximately double thrust magnitudes (22+/-6 mN, mean +/- S.D., N=20), resulting in a mean mechanical performance ratio (thrust/total force) of 0.32 at 1.2FL s(-1). An increase in speed by a factor of 1.8 resulted in a mean increase in thrust by a factor of 4.4, a mean increase in lateral forces by a factor of 3, no change in the magnitude of lift produced and an increase in mean mechanical performance to 0.42. The relatively high lateral forces generated during swimming may be a necessary consequence of force production via propagated waves of bending.

Animal Structures↗
PubMed2002

The C-start escape response of Polypterus senegalus: bilateral muscle activity and variation during stage 1 and 2.

The fast-start escape response is the primary reflexive escape mechanism in a wide phylogenetic range of fishes. To add detail to previously reported novel muscle activity patterns during the escape response of the bichir, Polypterus, we analyzed escape kinematics and muscle activity patterns in Polypterus senegalus using high-speed video and electromyography (EMG). Five fish were filmed at 250 Hz while synchronously recording white muscle activity at five sites on both sides of the body simultaneously (10 sites in total). Body wave speed and center of mass velocity, acceleration and curvature were calculated from digitized outlines. Six EMG variables per channel were also measured to characterize the motor pattern. P. senegalus shows a wide range of activity patterns, from very strong responses, in which the head often touched the tail, to very weak responses. This variation in strength is significantly correlated with the stimulus and is mechanically driven by changes in stage 1 muscle activity duration. Besides these changes in duration, the stage 1 muscle activity is unusual because it has strong bilateral activity, although the observed contralateral activity is significantly weaker and shorter in duration than ipsilateral activity. Bilateral activity may stiffen the body, but it does so by a constant amount over the variation we observed; therefore, P. senegalus does not modulate fast-start wave speed by changing body stiffness. Escape responses almost always have stage 2 contralateral muscle activity, often only in the anterior third of the body. The magnitude of the stage 2 activity is the primary predictor of final escape velocity.

Animals↗
PubMed2002

Quantification of the wake of rainbow trout (Oncorhynchus mykiss) using three-dimensional stereoscopic digital particle image velocimetry.

Although considerable progress has been made within the last decade in experimental hydrodynamic analyses of aquatic locomotion using two-dimensional digital particle image velocimetry (two-dimensional DPIV), data have been limited to simultaneous calculation of two out of the three flow velocity variables: downstream (U), vertical (V) and lateral (W). Here, we present the first biological application of stereo-DPIV, an engineering technique that allows simultaneous calculation of U, V and W velocity vectors. We quantified the wakes of rainbow trout (Oncorhynchus mykiss, 16.5-21.5 cm total body length, BL), swimming steadily in a recirculating flow tank at a slow cruising speed of 1.2 BL s(-1). These data extend the comparative basis of current hydromechanical data on the wakes of free-swimming fishes to the salmoniforms and are used to test current hypotheses of fin function by calculations of mechanical performance and Froude efficiency. Stereo-DPIV wake images showed three-dimensional views of oscillating jet flows high in velocity relative to free-stream values. These jet flows are consistent with the central momentum jet flows through the cores of shed vortex rings that have been previously viewed for caudal fin swimmers using two-dimensional DPIV. The magnitude and direction of U, V and W flows in these jets were determined over a time series of 6-8 consecutive strokes by each of four fish. Although the fish swam at the same relative speed, the absolute magnitudes of U, V and W were dependent on individual because of body size variation. Vertical flows were small in magnitude (<1 cm s(-1)) and variable in direction, indicating limited and variable vertical force production during slow, steady, forward swimming. Thus, in contrast to previous data from sunfish (Lepomis macrochirus) and mackerel (Scomber japonicus), the trout homocercal caudal fin does not appear to generate consistent vertical forces during steady swimming. U flows were of the order of 3-6 cm s(-1); lateral flows were typically strongest, with W magnitudes of 4-6 cm s(-1). Such strong lateral flows have also been shown for more derived euteleosts with homocercal caudal fins. The ratios of the magnitudes of wake flow, U/(U+V+W), which is a flow equivalent to mechanical performance, were also dependent on individual and ranged from 0.32 to 0.45, a range similar to the range of mechanical performance values previously determined using standard two-dimensional DPIV methods for caudal fin locomotion by more derived euteleosts. Strong lateral jet flow appears to be a general feature of caudal fin locomotion by teleosts and may reflect the nature of undulatory propulsion as a posteriorly propagated wave of bending. Froude efficiency (eta(p)) was independent of individual; mean eta(p) was 0.74, which is similar to previous findings for trout.

Animals↗