Living in a Turbulent World-A New Conceptual Framework for the Interactions of Fish and Eddies.

The natural habitats of fishes are characterized by movements of water driven by a multitude of physical processes of either natural or human origin. The resultant unsteadiness is exacerbated when flow interacts with surfaces, such as the bottom and banks, and protruding objects, such as corals, boulders, and woody debris. There is growing interest in the impacts on performance and behavior of fishes swimming in "turbulent flows". The ability of fishes to stabilize their postures and their swimming trajectories is thought to be important in determining species' distributions and densities, and hence the resultant assemblages in various habitats. A theoretical framework is proposed to quantify the interactions of fish and flows. Dimensionless parameters are derived based on a physical description of the flow structures and different regimes are predicted describing fishes' responses to a wide range of physical perturbations. We found the ratio of eddy size to fish size, the "momentum ratio" (ratio between momentum of the eddy and the momentum of the fish), as well as the time of interaction between eddy and fish to be especially important in determining thresholds for the fish's posture and trajectory.

Stability versus Maneuvering: Challenges for Stability during Swimming by Fishes.

Fishes are well known for their remarkable maneuverability and agility. Less visible is the continuous control of stability essential for the exploitation of the full range of aquatic resources. Perturbations to posture and trajectory arise from hydrostatic and hydrodynamic forces centered in a fish (intrinsic) and from the environment (extrinsic). Hydrostatic instabilities arise from vertical and horizontal separation of the centers of mass (CM) and of buoyancy, thereby creating perturbations in roll, yaw, and pitch, with largely neglected implications for behavioral ecology. Among various forms of hydrodynamic stability, the need for stability in the face of recoil forces from propulsors is close to universal. Destabilizing torques in body-caudal fin swimming is created by inertial and viscous forces through a propulsor beat. The recoil component is reduced, damped, and corrected in various ways, including kinematics, shape of the body and fins, and deployment of the fins. We postulate that control of the angle of orientation, θ, of the trailing edge is especially important in the evolution and lifestyles of fishes, but studies are few. Control of stability and maneuvering are reflected in accelerations around the CM. Accelerations for such motions may give insight into time-behavior patterns in the wild but cannot be used to determine the expenditure of energy by free-swimming fishes.

Assessing possible effects of fish-culture systems on fish swimming: the role of stability in turbulent flows.

Fish are cultured in ponds, recirculating systems, raceways, and cages. Turbulence is associated with one or more of mechanisms to facilitate food accessibility, maintain adequate levels of oxygen, remove carbon dioxide, urinary and fecal wastes, as well as from locomotion of fishes themselves. Turbulence has been shown to have positive and negative effects on fish swimming, feeding, and energetics, usually with negative impacts at very low and at high levels, and least effects and sometimes positive effects at intermediate levels. Differences in responses of fishes with varying levels of turbulence are related to the size of eddies relative to the size of a fish (larvae, juveniles, and adults). Impacts on locomotor functions are associated with eddy diameters of the order of 0.5-1L, where L is the total length of a fish. Negative locomotor impacts of turbulence are associated with eddies challenging stability, while positive effects promote drafting and station holding with reduced locomotor motions. Deployment of control surfaces increases with the level of turbulence up to a threshold where control is overwhelmed. The design of culture facilities is expected to affect levels of turbulence and may be engineered to provide optimal levels facilitating high growth.

Body-induced vortical flows: a common mechanism for self-corrective trimming control in boxfishes.

Boxfishes (Teleostei: Ostraciidae) are marine fishes having rigid carapaces that vary significantly among taxa in their shapes and structural ornamentation. We showed previously that the keels of the carapace of one species of tropical boxfish, the smooth trunkfish, produce leading edge vortices (LEVs) capable of generating self-correcting trimming forces during swimming. In this paper we show that other tropical boxfishes with different carapace shapes have similar capabilities. We conducted a quantitative study of flows around the carapaces of three morphologically distinct boxfishes (spotted boxfish, scrawled cowfish and buffalo trunkfish) using stereolithographic models and three separate but interrelated analytical approaches: digital particle image velocimetry (DPIV), pressure distribution measurements, and force balance measurements. The ventral keels of all three forms produced LEVs that grew in circulation along the bodies, resembling the LEVs produced around delta-winged aircraft. These spiral vortices formed above the keels and increased in circulation as pitch angle became more positive, and formed below the keels and increased in circulation as pitch angle became more negative. Vortices also formed along the eye ridges of all boxfishes. In the spotted boxfish, which is largely trapezoidal in cross section, consistent dorsal vortex growth posterior to the eye ridge was also present. When all three boxfishes were positioned at various yaw angles, regions of strongest concentrated vorticity formed in far-field locations of the carapace compared with near-field areas, and vortex circulation was greatest posterior to the center of mass. In general, regions of localized low pressure correlated well with regions of attached, concentrated vorticity, especially around the ventral keels. Although other features of the carapace also affect flow patterns and pressure distributions in different ways, the integrated effects of the flows were consistent for all forms: they produce trimming self-correcting forces, which we measured directly using the force balance. These data together with previous work on smooth trunkfish indicate that body-induced vortical flows are a common mechanism that is probably significant for trim control in all species of tropical boxfishes.

Investigation of an outbreak of encephalomyelitis caused by West Nile virus in 136 horses.

OBJECTIVE: To describe an outbreak of encephalomyelitis caused by West Nile virus (WNV) in horses in northern Indiana. DESIGN: Case series. ANIMALS: 170 horses. PROCEDURES: Horses with clinical signs suggestive of encephalomyelitis caused by WNV were examined. Date, age, sex, breed, and survival status were recorded. Serum samples were tested for anti-WNV antibodies, and virus isolation was attempted from samples of brain tissue. Climate data from local weather recording stations were collected. An epidemic curve was constructed, and case fatality rate was calculated. RESULTS: The most common clinical signs were ataxia, hind limb paresis, and muscle tremors and fasciculations. Eight horses had been vaccinated against WNV from 2 to 21 days prior to the appearance of clinical signs. West Nile virus was isolated from brain tissue of 2 nonvaccinated horses, and anti-WNV IgM antibodies were detected in 132 nonvaccinated horses; in 2 other nonvaccinated horses, anti-WNV antibodies were detected and WNV was also isolated from brain tissue. Thirty-one (22.8%) horses died or were euthanatized. The peak of the outbreak occurred on September 6, 2002. Ambient temperatures were significantly lower after the peak of the outbreak, compared with prior to the peak. CONCLUSIONS AND CLINICAL RELEVANCE: The peak risk period for encephalomyelitis caused by WNV in northern Indiana was mid-August to mid-September. Reduction in cases coincided with decreasing ambient temperatures. Because of a substantial case fatality rate, owners of horses in northern Indiana should have their horses fully protected by vaccination against WNV before June. In other regions of the United States with a defined mosquito breeding season, vaccination of previously nonvaccinated horses should commence at least 4 months before the anticipated peak in seasonal mosquito numbers, and for previously vaccinated horses, vaccine should be administered no later than 2 months before this time.

Response latencies to postural disturbances in three species of teleostean fishes.

Flow in aquatic systems is characterized by unsteadiness that creates destabilizing perturbations. Appropriate correction responses depend on response latency. The time between a disturbance induced by either removal of a flow refuge or striking various parts of the body with a narrow water jet was measured for three species, chosen as examples of modes in teleostean body/fin organization that are expected to affect stability. Creek chub Semotilus atromaculatus is representative of fusiform-bodied soft-rayed teleosts, smallmouth bass Micropterus dolomieu of fusiform-bodied spiny-rayed forms and bluegill Lepomis macrochirus of deep-bodied spiny-rayed forms. Observations were made at 23 degrees C. Loss of refuge resulted in a surge that fish corrected by starting to swim within 129+/-29 ms (mean +/- 2 S.E.M.) for chub, which was significantly shorter than minimal times of approximately 200 ms for bluegill and bass. Slips and heaves induced by water jets initially resulted in extension of the median and paired fins that would damp growth of the disturbance, but otherwise these disturbances were ignored. Yaws and pitches were more likely to cause fish to swim away from the stimulus, making corrections as they did so. There were no differences in latencies for slip, heave, yaw and pitch disturbances within each species, but latencies varied among species. For these disturbances, responses averaged 123+/-19 ms for chub, again significantly smaller than those of 201+/-24 ms for bass and 208+/-52 ms for bluegill. Values for the two centrarchids were not significantly different (P>0.08). The response latency for rolling disturbances did not differ among species but was significantly smaller than that for other disturbances, with an overall latency of 70+/-15 ms. The greater responsiveness to hydrostatic rolling instability is attributed to functions requiring an upright posture and differences among species in habitat preferences.

Hydrodynamic stability of swimming in ostraciid fishes: role of the carapace in the smooth trunkfish Lactophrys triqueter (Teleostei: Ostraciidae).

The hydrodynamic bases for the stability of locomotory motions in fishes are poorly understood, even for those fishes, such as the rigid-bodied smooth trunkfish Lactophrys triqueter, that exhibit unusually small amplitude recoil movements during rectilinear swimming. We have studied the role played by the bony carapace of the smooth trunkfish in generating trimming forces that self-correct for instabilities. The flow patterns, forces and moments on and around anatomically exact, smooth trunkfish models positioned at both pitching and yawing angles of attack were investigated using three methods: digital particle image velocimetry (DPIV), pressure distribution measurements, and force balance measurements. Models positioned at various pitching angles of attack within a flow tunnel produced well-developed counter-rotating vortices along the ventro-lateral keels. The vortices developed first at the anterior edges of the ventro-lateral keels, grew posteriorly along the carapace, and reached maximum circulation at the posterior edge of the carapace. The vortical flow increased in strength as pitching angles of attack deviated from 0 degrees, and was located above the keels at positive angles of attack and below them at negative angles of attack. Variation of yawing angles of attack resulted in prominent dorsal and ventral vortices developing at far-field locations of the carapace; far-field vortices intensified posteriorly and as angles of attack deviated from 0 degrees. Pressure distribution results were consistent with the DPIV findings, with areas of low pressure correlating well with regions of attached, concentrated vorticity. Lift coefficients of boxfish models were similar to lift coefficients of delta wings, devices that also generate lift through vortex generation. Furthermore, nose-down and nose-up pitching moments about the center of mass were detected at positive and negative pitching angles of attack, respectively. The three complementary experimental approaches all indicate that the carapace of the smooth trunkfish effectively generates self-correcting forces for pitching and yawing motions--a characteristic that is advantageous for the highly variable velocity fields experienced by trunkfish in their complex aquatic environment. All important morphological features of the carapace contribute to producing the hydrodynamic stability of swimming trajectories in this species.

Kinematics of plaice, Pleuronectes platessa, and cod, Gadus morhua, swimming near the bottom.

The kinematics of plaice (Pleuronectes platessa, L=22.1 cm) and cod (Gadus morhua, L=25.0 cm, where L is total fish length) swimming at various speeds at the bottom and lifted to heights, h, of 10, 50 and 100 mm by a thin-wire grid were measured. For cod, tailbeat frequency, amplitude, body and fin span and propulsive wavelength were unaffected by h and varied with speed as described for fusiform pelagic species. In contrast, the kinematics of plaice was affected by h. Body and fin spans and propulsive wavelength were independent of swimming speed and h. Tailbeat amplitude was independent of swimming speed, but averaged 1.5 cm at h=0 and 2.5 cm at h> or = 10 mm. Plaice tailbeat frequency increased with swimming speed for fish at the bottom but was independent of swimming speed at h=10, 50 and 100 mm, averaging 4.6, 6.0 and 5.8 Hz respectively. Total mechanical power, P, produced by propulsive movements calculated from the bulk-momentum form of elongated slender-body theory was similar for cod and plaice swimming at the bottom but, at h> or = 10 mm, P for plaice was larger than that for cod. Plaice support their weight in water by swimming at a small tilt angle. The small changes in swimming kinematics with swimming speed are attributed to decreasing induced power costs to support the weight as speed increases. The contribution of the tail to power output increased monotonically with the tail gap/span ratio, z/B, for z/B=0.23 (h=0 mm) to z/B=1.1 (h=50 mm). The smaller tailbeat amplitude of the tail decreased both z/B and the power output for plaice swimming at the bottom. For the maximum body and fin span of plaice, the contribution to power output increased for local z/B values of 0.044 (h-0 mm) to 0.1 (h=10 mm) and declined somewhat at larger values of z/B. The smaller effect of the bottom on power output of the large-span anterior body sections may result from the resorption of much of the upstream wake at the re-entrant downstream tail.

Flow Patterns Around the Carapaces of Rigid-bodied, Multi-propulsor Boxfishes (Teleostei: Ostraciidae).

Boxfishes (Teleostei: Ostraciidae) are rigid-body, multi-propulsor swimmers that exhibit unusually small amplitude recoil movements during rectilinear locomotion. Mechanisms producing the smooth swimming trajectories of these fishes are unknown, however. Therefore, we have studied the roles the bony carapaces of these fishes play in generating this dynamic stability. Features of the carapaces of four morphologically distinct species of boxfishes were measured, and anatomically-exact stereolithographic models of the boxfishes were constructed. Flow patterns around each model were investigated using three methods: 1) digital particle image velocimetry (DPIV), 2) pressure distribution measurements, and 3) force balance measurements. Significant differences in both cross-sectional and longitudinal carapace morphology were detected among the four species. However, results from the three interrelated approaches indicate that flow patterns around the various carapaces are remarkably similar. DPIV results revealed that the keels of all boxfishes generate strong longitudinal vortices that vary in strength and position with angle of attack. In areas where attached, concentrated vorticity was detected using DPIV, low pressure also was detected at the carapace surface using pressure sensors. Predictions of the effects of both observed vortical flow patterns and pressure distributions on the carapace were consistent with actual forces and moments measured using the force balance. Most notably, the three complementary experimental approaches consistently indicate that the ventral keels of all boxfishes, and in some species the dorsal keels as well, effectively generate self-correcting forces for pitching motions-a characteristic that is advantageous for the highly variable velocity fields in which these fishes reside.

Power requirements of swimming: do new methods resolve old questions?

A recurring question in the study of fish biomechanics and energetics is the mechanical power required for tail-swimming at the high speeds seen among aquatic vertebrates. The quest for answers has been driven by conceptual advances in fluid dynamics, starting with ideas on the boundary layer and drag initiated by Prandtl, and in measurement techniques starting with force balances focussing on drag and thrust. Drag (=thrust) from measurements on physical models, carcasses, kinematics as inputs to hydromechanical models, and physiological power sources vary from less than that expected for an equivalent rigid reference to over an order of magnitude greater. Estimates of drag and thrust using recent advances largely made possible by increased computing power have not resolved the discrepancy. Sources of drag and thrust are not separable in axial undulatory self propulsion, are open to interpretation and Froude efficiency is zero. Wakes are not easily interpreted, especially for thrust evaluation. We suggest the best measures of swimming performance are velocity and power consumption for which 2D inviscid simulations can give realistic predictions. Steady swimming power is several times that required for towing an equivalent flat plate at the same speed.

Control of posture, depth, and swimming trajectories of fishes.

Perturbations vary in period and amplitude, and responses to unavoidable perturbations depend on response time and scale. Disturbances due to unavoidable perturbations occur in three translational planes and three rotational axes during forwards and backwards swimming. Stability depends on hydrodynamic damping and correcting forces, which may be generated by propulsors (powered) or by control surfaces moving with the body (trimming). Hydrostatic forces affecting body orientation (posture) result in negative metacentric heights amplifying rolling disturbances. The ability to counteract perturbations and correct disturbances is greater for fishes with more slender bodies, which appears to affect habitat choices. Postural control problems are greatest at low speeds, and are avoided by some fishes by sitting on the bottom. In currents, body form and behavior affect lift, drag, weight, and friction and hence speeds to which posture can be controlled. Self-correcting and regulated damping and trimming mechanisms are most important in stabilizing swimming trajectories. Body resistance, fin trajectory, multiple propulsors, and long-based fins damp self-generated locomotor disturbances. Powered control using the tail evolved early in chordates, and is retained by most groups, although fishes, especially acanthopterygians, make greater use of appendages. As with most areas of stability, little is known of control costs. Costs and benefits of low-density inclusions and hydrodynamic mechanisms for depth control vary with habits and habitats. Control may make substantial contributions to energy budgets.