Function of the Heterocercal Tail in Sharks: Quantitative Wake Dynamics During Steady Horizontal Swimming and Vertical Maneuvering

The function of the heterocercal tail in sharks has long been debated in the literature. Previous kinematic data have supported the classical theory which proposes that the beating of the heterocercal caudal fin during steady horizontal locomotion pushes posteroventrally on the water, generating a reactive force directed anterodorsally and causing rotation around the center of mass. An alternative model suggests that the heterocercal shark tail functions to direct reaction forces through the center of mass. In this paper, we quantify the function of the tail in two species of shark and compare shark tail function with previous hydrodynamic data on the heterocercal tail of sturgeon Acipenser transmontanus. To address the two models of shark heterocercal tail function, we applied the technique of digital particle image velocimetry (DPIV) to quantify the wake of two species of shark swimming in a flow tank. Both steady horizontal locomotion and vertical maneuvering were analyzed. We used DPIV with both horizontal and vertical light sheet orientations to quantify patterns of wake velocity and vorticity behind the heterocercal tail of leopard sharks (Triakis semifasciata) and bamboo sharks (Chiloscyllium punctatum) swimming at 1.0 Ls–1, where L is total body length. Two synchronized high-speed video cameras allowed simultaneous measurement of shark body position and wake structure. We measured the orientation of tail vortices shed into the wake and the orientation of the central jet through the core of these vortices relative to body orientation. Analysis of flow geometry indicates that the tail of both leopard and bamboo shark generates strongly tilted vortex rings with a mean jet angle of approximately 30 ° below horizontal during steady horizontal swimming. The corresponding angle of the reaction force is much greater than body angle (mean 11 °) and the angle of the path of motion of the center of mass (mean approximately 0 °), thus strongly supporting the classical model of heterocercal tail function for steady horizontal locomotion. Vortex jet angle varies significantly with body angle changes during vertical maneuvering, but sharks show no evidence of active reorientation of jet angle relative to body angle, as was seen in a previous study on the function of sturgeon tail. Vortex jet orientation is significantly more inclined than the relatively horizontal jet generated by sturgeon tail vortex rings, demonstrating substantial differences in function in the heterocercal tails of sharks and sturgeon. We present a summary of forces on a swimming shark integrating data obtained here on the tail with previous data on pectoral fin and body function. Body orientation plays a critical role in the overall force balance and compensates for torques generated by the tail. The pectoral fins do not generate lift during steady horizontal locomotion, but play an important hydrodynamic role during vertical maneuvering

On the rules for aquatic locomotion

We present unifying rules governing the efficient locomotion of swimming fish and marine mammals. Using scaling and dimensional analysis, supported by new experimental data, we show that efficient locomotion occurs when the values of the Strouhal (St) number St(=f A/U) and A∗(=A/L), two nondimensional numbers that relate forward speed U, tail-beat amplitude A, tail-beat frequency f , and the length of the swimmer L are bound to the tight ranges of 0.2–0.4 and 0.1–0.3, respectively. The tight range of 0.2–0.4 for the St number has previously been associated with optimal thrust generation. We show that the St number alone is insufficient to achieve optimal aquatic locomotion, and an additional condition on A∗ is needed. More importantly, we show that when swimming at minimal power consumption, the Strouhal number of a cruising swimmer is predetermined solely by the shape and drag characteristics of the swimmer. We show that diverse species of fish and cetaceans cruise indeed with the St number and A∗ predicted by our theory. Our findings provide a physical explanation as to why fast aquatic swimmers cruise with a relatively constant tail-beat amplitude of approximately 20% of the body length, and their swimming speed is nearly proportional to their tail-beat frequenc

Microwave Current Imaging in Passive HTS Components by Low-Temperature Laser Scanning Microscopy (LTLSM)

We have used the LTLSM technique for a spatially resolved investigation of the microwave transport properties, nonlinearities and material inhomogeneities in an operating coplanar waveguide YBa_2Cu_3O_{7-\delta} (YBCO) microwave resonator on an LaAlO_3 (LAO) substrate. The influence of twin-domain blocks, in-plane rotated grains, and micro-cracks in the YBCO film on the nonuniform rf current distribution were measured with a micrometer-scale spatial resolution. The impact of the peaked edge currents and rf field penetration into weak links on the linear device performance were studied as well. The LTLSM capabilities and its future potential for non-destructive characterization of the microwave properties of superconducting circuits are discussed.Comment: 8 pages, 9 figures, 2-column format, presented at High Temperature Superconductors in High Frequency Fields 2004, Journal of Superconductivity (in press

Fish optimize sensing and respiration during undulatory swimming

Previous work in fishes considers undulation as a means of propulsion without addressing how it may affect other functions such as sensing and respiration. Here we show that undulation can optimize propulsion, flow sensing and respiration concurrently without any apparent tradeoffs when head movements are coupled correctly with the movements of the body. This finding challenges a long-held assumption that head movements are simply an unintended consequence of undulation, existing only because of the recoil of an oscillating tail. We use a combination of theoretical, biological and physical experiments to reveal the hydrodynamic mechanisms underlying this concerted optimization. Based on our results we develop a parsimonious control architecture that can be used by both undulatory animals and machines in dynamic environments

The development of a biologically inspired propulsor for unmanned underwater vehicles

IEEE Journal of Oceanic Engineering, 32(3): pp. 533-550Fish are remarkable in their ability to maneuver and to control their body position. This ability is the result of the coordinated movement of fins which extend from the body and form control surfaces that can create and vector forces in 3-D. We have embarked on a research program designed to develop a maneuvering propulsor for unmanned undersea vehicles (UUVs) that is based on the pectoral fin of the bluegill sunfish. For this, the anatomy, kinematics, and hydrodynamics of the sunfish pectoral fin were investigated experimentally and through the use of computational fluid dynamics (CFD) simulations. These studies identified that the kinematics of the sunfish pectoral fin are very complex and are not easily described by traditional “rowing”- and “flapping”-type kinematics. A consequence of the complex motion is that the pectoral fin can produce forward thrust during both its outstroke (abduction) and instroke (adduction), and while doing so generates only small lateral and lift forces. The results of the biological studies were used to guide the design of robotic pectoral fins which were built as experimental devices and used to investigate the mechanisms of thrust production and control. Because of a design that was based heavily on the anatomy of the sunfish fin, the robotic pectoral fins had the level of control and degrees of freedom necessary to reproduce many of the complex fin motions used by the sunfish during steady swimming. These robotic fins are excellent experimental tools, and are an important first step towards developing propulsive devices that will give the next generation of UUVs the ability to produce and control thrust like highly maneuverable fish

The Science of Phylogenetic Systematics: Explanation, Prediction, and Test

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/73926/1/j.1096-0031.1999.tb00279.x.pd

Neurotrophic actions of dopamine on the development of a serotonergic feeding circuit in Drosophila melanogaster

<p>Abstract</p> <p>Background</p> <p>In the fruit fly, <it>Drosophila melanogaster</it>, serotonin functions both as a neurotransmitter to regulate larval feeding, and in the development of the stomatogastric feeding circuit. There is an inverse relationship between neuronal serotonin levels during late embryogenesis and the complexity of the serotonergic fibers projecting from the larval brain to the foregut, which correlate with perturbations in feeding, the functional output of the circuit. Dopamine does not modulate larval feeding, and dopaminergic fibers do not innervate the larval foregut. Since dopamine can function in central nervous system development, separate from its role as a neurotransmitter, the role of neuronal dopamine was assessed on the development, and mature function, of the 5-HT larval feeding circuit.</p> <p>Results</p> <p>Both decreased and increased neuronal dopamine levels in late embryogenesis during development of this circuit result in depressed levels of larval feeding. Perturbations in neuronal dopamine during this developmental period also result in greater branch complexity of the serotonergic fibers innervating the gut, as well as increased size and number of the serotonin-containing vesicles along the neurite length. This neurotrophic action for dopamine is modulated by the D₂dopamine receptor expressed during late embryogenesis in central 5-HT neurons. Animals carrying transgenic RNAi constructs to knock down both dopamine and serotonin synthesis in the central nervous system display normal feeding and fiber architecture. However, disparate levels of neuronal dopamine and serotonin during development of the circuit result in abnormal gut fiber architecture and feeding behavior.</p> <p>Conclusions</p> <p>These results suggest that dopamine can exert a direct trophic influence on the development of a specific neural circuit, and that dopamine and serotonin may interact with each other to generate the neural architecture necessary for normal function of the circuit.</p

Development of mandibular, hyoid and hypobranchial muscles in the zebrafish: homologies and evolution of these muscles within bony fishes and tetrapods

<p>Abstract</p> <p>Background</p> <p>During vertebrate head evolution, muscle changes accompanied radical modification of the skeleton. Recent studies have suggested that muscles and their innervation evolve less rapidly than cartilage. The freshwater teleostean zebrafish (<it>Danio rerio</it>) is the most studied actinopterygian model organism, and is sometimes taken to represent osteichthyans as a whole, which include bony fishes and tetrapods. Most work concerning zebrafish cranial muscles has focused on larval stages. We set out to describe the later development of zebrafish head muscles and compare muscle homologies across the Osteichthyes.</p> <p>Results</p> <p>We describe one new muscle and show that the number of mandibular, hyoid and hypobranchial muscles found in four day-old zebrafish larvae is similar to that found in the adult. However, the overall configuration and/or the number of divisions of these muscles change during development. For example, the undivided adductor mandibulae of early larvae gives rise to the adductor mandibulae sections A0, A1-OST, A2 and Aω, and the protractor hyoideus becomes divided into dorsal and ventral portions in adults. There is not always a correspondence between the ontogeny of these muscles in the zebrafish and their evolution within the Osteichthyes. All of the 13 mandibular, hyoid and hypobranchial muscles present in the adult zebrafish are found in at least some other living teleosts, and all except the protractor hyoideus are found in at least some extant non-teleost actinopterygians. Of these muscles, about a quarter (intermandibularis anterior, adductor mandibulae, sternohyoideus) are found in at least some living tetrapods, and a further quarter (levator arcus palatini, adductor arcus palatini, adductor operculi) in at least some extant sarcopterygian fish.</p> <p>Conclusion</p> <p>Although the zebrafish occupies a rather derived phylogenetic position within actinopterygians and even within teleosts, with respect to the mandibular, hyoid and hypobranchial muscles it seems justified to consider it an appropriate representative of these two groups. Among these muscles, the three with clear homologues in tetrapods and the further three identified in sarcopterygian fish are particularly appropriate for comparisons of results between the actinopterygian zebrafish and the sarcopterygians.</p

Sophisticated falsification and research cycles: Consequences for differential character weighting in phylogenetic systematics

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/72703/1/j.1463-6409.1997.tb00424.x.pd