Numerical Study of Propulsion Mechanism for Oscillating Rigid and Flexible Tuna-Tails

Numerical study on the unsteady hydrodynamic characteristics of oscillating rigid and flexible tuna-tails in viscous flow-field is performed.Investigations are conducted using Reynolds-Averaged Navier-Stokes (RANS) equations with a moving adaptive mesh.The effect of swimming speed,flapping amplitude,frequency and flexure amplitude on the propulsion performance of the rigid and flexible tuna-tails are investigated.Computational results reveal that a pair of leading edge vortices develop along the tail surface as it undergoes an oscillating motion.The propulsive efficiency has a strong correlation with various locomotive parameters.Peak propulsive efficiency can be obtained by adjusting these parameters.Particularly,when input power coefficient is less than 2.8,the rigid tail generates larger thrust force and higher propulsive efficiency than flexible tail.However,when input power coefficient is larger than 2.8,flexible tail is superior to rigid tail.     

Effect of Chord Flexure on Aerodynamic Performance of a Flapping Wing

Inspired by the fact that a high flexible wing in nature generates high aerodynamic performance, we investigated the aerodynamic performance of the flapping wing with different chord flexures. The unsteady, incompressible, and viscous flow over airfoil NACA0012 in a plunge motion was analyzed by using Navier-Stokes equation. Grid deformation, in which finite element and interpolation ideas are mixed, was introduced for computing large grid deformation caused by the chord flexures. We explored the optimal phase angle for thrust force and propulsive efficiency by varying the chord flexure from 0.05 to 0.7 when reduced frequency and plunge amplitude were fixed. Throughout parametric study on the phase angle and chord flexure amplitude, the maximum thrust force is achieved near at 0° in all given conditions, meanwhile, it is found that the optimal phase angle has dependency of chord flexure amplitude, which achieves higher aerodynamic performance compared to previous studies. These findings will provide a useful guideline for determining wing flexibility in design of a bio-mimetic air vehicle.     

Foreflipper propulsion in the California sea lion, Zalophus californianus

The family Otariidae comprises the only group of marine mammals that habitually use their pectoral appendages to generate propulsive forces during swimming. This method of propulsion was examined in the California sea lion ( Zalophus californianus ), a representative member of the family. High-speed films were taken as a sea lion swam against a water current generated inside a large flow channel. Thrust production was determined by examining the body's movement at various stages of the propulsive cycle. Sea lions generate thrust continuously throughout the stroke. Over its initial three-quarters, foreflippers act as hydrofoils creating forward thrust and lift as they move vertically through the water. Thrust production is greatest, however, near the end of the stroke, when flippers are used as paddles and are oriented broad side to the oncoming flow. The force generated by this three-phased system of propulsion is likely to be greater than that attainable by either an exclusively lift-based hydrofoil or drag-based paddling style of swimming.
The kinematic changes that enable sea lions to change speed were also investigated. Film records revealed that stroke amplitude became greater with speed, although total stroke duration remained essentially constant. Sea lions increase stroke frequency with velocity but large variations in the measured values suggest that changes in amplitude and flipper angle of attack are also important parameters for modulating swimming speed.     

Morphological adaptations to marine life in snakes

We investigated morphological adaptations to aquatic life within animals that exhibit a structurally simple, elongate body form, i.e., snakes. This linear body plan should impose different biomechanical constraints than the classical streamlined body shape associated with propulsion by fins, feet, or wings. Our measurements of general body shape of terrestrial, amphibious, and marine snakes (all from the same phylogenetic lineage, the Elapidae) show that seasnakes display specialized morphological attributes for life in water. Most notably, the cross‐sectional body shape is circular in terrestrial snakes but dorso‐ventrally elongated in seasnakes (due to a prominent ventral keel); amphibious species (sea kraits) exhibit an intermediate shape. The tail of amphibious and marine species (a major propulsive structure during swimming) is higher and thinner than in terrestrial snakes (i.e., paddle‐shaped) but shorter relative to body length. The evolution of a laterally compressed shape has been achieved by an increase in body height rather than a decrease in body width, possibly reflecting selection for more effective propulsive thrust, and for an ability to maintain hydrodynamic efficiency despite the minor bodily distension inevitably caused by prey items and developing offspring. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc     

Advantages of a Biomimetic Stiffness Profile in Pitching Flexible Fin Propulsion

<正> The use of oscillating flexible fins in propulsion has been the subject of several studies in recent years, but attention israrely paid to the specific role of stiffness profile in thrust production.Stiffness profile is defined as the variation in localchordwise bending stiffness (EI) of a fin, from leading to trailing edge.In this study, flexible fins with a standard NACA0012shape were tested alongside fins with a stiffness profile mimicking that of a Pumpkinseed Sunfish (Lepomis gibbosus).The finswere oscillated with a pitching sinusoidal motion over a range of frequencies and amplitudes, while torque, lateral force andstatic thrust were measured.Over the range of oscillation parameters tested, it was shown that the fin with a biomimetic stiffness profile offered a significantimprovement in static thrust, compared to a fin of similar dimensions with a standard NACA0012 aerofoil profile.Thebiomimetic fin also produced thrust more consistently over each oscillation cycle.A comparison of fin materials of different stiffness showed that the improvement was due to the stiffness profile itself, andwas not simply an effect of altering the overall stiffness of the fin.Fins of the same stiffness profile were observed to follow thesame thrust-power curve, independent of the stiffness of the moulding material.Biomimetic fins were shown to produce up to26% greater thrust per watt of input power, within the experimental range.     

Separation of propulsive and adhesive traction stresses in locomoting keratocytes

Strong, actomyosin-dependent, pinching tractions in steadily locomoting (gliding) fish keratocytes revealed by traction imaging present a paradox, since only forces perpendicular to the direction of locomotion are apparent, leaving the actual propulsive forces unresolved. When keratocytes become transiently "stuck" by their trailing edge and adopt a fibroblast-like morphology, the tractions opposing locomotion are concentrated into the tail, leaving the active pinching and propulsive tractions clearly visible under the cell body. Stuck keratocytes can develop approximately 1 mdyn (10,000 pN) total propulsive thrust, originating in the wings of the cell. The leading lamella develops no detectable propulsive traction, even when the cell pulls on its transient tail anchorage. The separation of propulsive and adhesive tractions in the stuck phenotype leads to a mechanically consistent hypothesis that resolves the traction paradox for gliding keratocytes: the propulsive tractions driving locomotion are normally canceled by adhesive tractions resisting locomotion, leaving only the pinching tractions as a resultant. The resolution of the traction pattern into its components specifies conditions to be met for models of cytoskeletal force production, such as the dynamic network contraction model (Svitkina, T.M., A.B. Verkhovsky, K.M. McQuade, and G.G. Borisy. 1997. J. Cell Biol. 139:397-415). The traction pattern associated with cells undergoing sharp turns differs markedly from the normal pinching traction pattern, and can be accounted for by postulating an asymmetry in contractile activity of the opposed lateral wings of the cell.     

Experimental and Numerical Study of Penguin Mode Flapping Foil Propulsion System for Ships

The use of biomimetic tandem flapping foils for ships and underwater vehicles is considered as a unique and interesting concept in the area of marine propulsion.The flapping wings can be used as a thrust producing,stabilizer and control devices which has both propulsion and maneuvering applications for marine vehicles.In the present study,the hydrodynamic performance of a pair of flexible flapping foils resembling penguin flippers is studied.A ship model of 3 m in length is fitted with a pair of counter flapping foils at its bottom mid-ship region.Model tests are carried out in a towing tank to estimate the propulsive performance of flapping foils in bollard and self propulsion modes.The same tests are performed in a numerical environment using a Computational Fluid Dynamics (CFD) software.The numerical and experimental results show reasonably good agreement in both bollard pull and self propulsion trials.The numerical studies are carried out on flexible flapping hydrofoil in unsteady conditions using moving unstructured grids.The efficiency and force coefficients of the flexible flapping foils are determined and presented as a function of Strouhal number.     

Hydrodynamic design of the humpback whale flipper

The humpback whale (Megaptera novaeangliae) is reported to use its elongate pectoral flippers during swimming maneuvers. The morphology of the flipper from a 9.02-m whale was evaluated with regard to this hydrodynamic function. The flipper had a wing-like, high aspect ratio plan-form. Rounded tubercles were regularly interspersed along the flipper's leading edge. The flipper was cut into 71 2.5-cm cross-sections and photographed. Except for sections near the distal tip, flipper sections were symmetrical with no camber. Flipper sections had a blunt, rounded leading edge and a highly tapered trailing edge. Placement of the maximum thickness placement for each cross-section varied from 49% of chord at the tip to 19% at mid-span. Section thickness ratio averaged 0.23 with a range of 0.20–0.28. The humpback whale flipper had a cross-sectional design typical of manufactured aerodynamic foils for lift generation. The morphology and placement of leading edge tubercles sugges that they function as enhanced lift devices to control flow over the flipper and maintain lift at high angles of attack. The morphology of the humpback whale flipper suggests that it is adapted for high maneuverability associated with the whale's unique feeding behavior. © 1995 Wiley-Liss, Inc.     

Bridging “Romer’s Gap”: Limb Mechanics of an Extant Belly-Dragging Lizard Inform Debate on Tetrapod Locomotion During the Early Carboniferous

Devonian stem tetrapods are thought to have used ‘crutching’ on land, a belly-dragging form of synchronous forelimb action-powered locomotion. During the Early Carboniferous, early tetrapods underwent rapid radiation, and the terrestrial locomotion of crown-group node tetrapods is believed to have been hindlimb-powered and ‘raised’, involving symmetrical gaits similar to those used by modern salamanders. The fossil record over this period of evolutionary transition is remarkably poor (Romer’s Gap), but we hypothesize a phase of belly-dragging sprawling locomotion combined with symmetrical gaits. Since belly-dragging sprawling locomotion has differing functional demands from ‘raised’ sprawling locomotion, we studied the limb mechanics of the extant belly-dragging blue-tongued skink. We used X-ray reconstruction of moving morphology to quantify the three-dimensional kinematic components, and simultaneously recorded single limb substrate reaction forces (SRF) in order to calculate SRF moment arms and the external moments acting on the proximal limb joints. In the hindlimbs, stylopodal long-axis rotation is more emphasized than in the forelimbs, and much greater vertical and propulsive forces are exerted. The SRF moment arm acting on the shoulder is at a local minimum at the instant of peak force. The hindlimbs display patterns that more closely resemble ‘raised’ sprawling species. External moment at the shoulder of the skink is smaller than in ‘raised’ sprawlers. We propose an evolutionary scenario in which the locomotor mechanics of belly-dragging early tetrapods were gradually modified towards hindlimb-powered, raised terrestrial locomotion with symmetrical gait. In accordance with the view that limb evolution was an exaptation for terrestrial locomotion, the kinematic pattern of the limbs for the generation of propulsion preceded, in our scenario, the evolution of permanent body weight support.     

Patterns of ossification in the manus of the harbor porpoise (Phocoena phocoena): hyperphalangy and delta-shaped bones

With the transition from terrestrial to aquatic habitats, cetacean forelimbs have undergone significant modifications in bone morphology and soft tissue distribution. Some, but not all, of these modifications are also demonstrated in other lineages of extant and extinct secondarily aquatic tetrapods. This study examines the ontogenetic pattern of ossification of the manus of the harbor porpoise (Phocoena phocoena), using plain film radiography. Two modifications examined are hyperphalangy (number of phalanges per digit increased beyond the typical mammalian number) and the morphology of delta-shaped bones. Hyperphalangy in Phocoena phocoena is apparent in digits 2 and 3. Phalangeal counts in all digits are variable (sometimes between the right and left flippers of the same individual) and are not necessarily correlated with age. Phalangeal ossification and epiphyseal fusion proceeds along the proximo-distal axis within each digit. In addition, digits 2 and 3 are at a more advanced stage of ossification than more abaxial digits. Delta-shaped bones appear to be a normal stage in the ossification of phalanges in all digits except the third, and may persist in the adult in certain digits. In humans, this morphology is a developmental anomaly usually associated with other malformations, such as polydactyly or syndactyly. Delta-shaped bones in the cetacean manus display a consistent orientation and the process by which they are formed may be similar to that in extinct marine reptiles.     

Functional differentiation of trailing and leading forelimbs during locomotion on the ground and on a horizontal branch in the European red squirrel (Sciurus vulgaris, Rodentia)

Mammalian locomotion is characterized by the frequent use of in-phase gaits in which the footfalls of the left and right fore- or hindlimbs are unevenly spaced in time. Although previous studies have identified a functional differentiation between the first limb (trailing limb) and the second limb (leading limb) to touch the ground during terrestrial locomotion, the influence of a horizontal branch on limb function has never been explored. To determine the functional differences between trailing and leading forelimbs during locomotion on the ground and on a horizontal branch, X-ray motion analysis and force measurements were carried out in two European red squirrels (Sciurus vulgaris, Rodentia). The differences observed between trailing and leading forelimbs were minimal during terrestrial locomotion, where both limbs fulfill two functions and go through a shock-absorbing phase followed by a generating phase. During locomotion on a horizontal branch, European red squirrels reduce speed and all substrate reaction forces transmitted may be due to the reduction of vertical oscillation of the center of mass. Further adjustments during locomotion on a horizontal branch differ significantly between trailing and leading forelimbs and include limb flexion, lead intervals, limb protraction and vertical displacement of the scapular pivot. Consequently, trailing and leading forelimbs perform different functions. Trailing forelimbs function primarily as shock-absorbing elements, whereas leading forelimbs are characterized by a high level of stiffness. This functional differentiation indicates that European red squirrels ‘test’ the substrate for stability with the trailing forelimb, while the leading forelimb responds to or counteracts swinging or snapping branches.     

Evolutionary and structural diversification of the larval nervous system among marine bryozoans

Regardless of the morphological divergence among larval forms of marine bryozoans, the larval nervous system and its major effector organs (musculature and ciliary fields) are largely molded on the basis of functional demands of feeding, ciliary propulsion, phototactic behaviors, and substrate exploration. Previously published ultrastructural information and immunohistochemical reconstructions presented here indicate that neuronal pathways are largely ipsilateral, with more complex synaptic connections localized within the nerve nodule. Multiciliated sensory-motor neurons diversify structurally and functionally on the basis of their position along the axis of swimming largely due to the functional demands of photoklinotaxis and substrate exploration. Vesiculariform, buguliform, and ascophoran coronate larvae all have patches of sensory neurons bordering the pyriform organ's ciliated groove (juxtapapillary cells and border cells) that are active during substrate selection. Despite their simplified form, cyclostome larvae maintain swimming and probing behaviors with sensory-motor systems functionally similar to those of some parenchymella and planula larval types. Considering the evolutionary relationships among the morphological grades of marine bryozoans, particular lineages within the gymnolaemates have independently evolved larval traits that convey a greater range of sensory abilities and increased propulsive capacity. The larval nervous system of bryozoans may be evolutionarily derived from the pretrochal region of a trochophore-like larval form.     

Independent innovation in the evolution of paddle-shaped tails in viviparous sea snakes (Elapidae: Hydrophiinae)

The viviparous sea snakes (Hydrophiinae) comprise ~90% of living marine reptiles and display many physical and behavioral adaptations for breathing, diving, and achieving osmotic balance in marine habitats. Among the most important innovations found in marine snakes are their paddle-shaped (dorsoventrally expanded) tails, which provide propulsive thrust in the dense aquatic medium. Here, we reconstruct the evolution of caudal paddles in viviparous sea snakes using a dated molecular phylogeny for all major lineages and computed tomography of internal osteological structures. Bayesian ancestral state reconstructions show that extremely large caudal paddles supported by elongated vertebral processes are unlikely to have been present in the most recent common ancestor of extant sea snakes. Instead, these characters appear to have been acquired independently in two highly marine lineages of relatively recent origin. Both the Aipysurus and Hydrophis lineages have elongated neural spines that support the dorsal edge of their large paddles. However, whereas in the Aipysurus lineage the ventral edge of the paddle is supported by elongated haemapophyses, this support is provided by elongated and ventrally directed pleurapophyses in the Hydrophis lineage. Three semi-marine lineages (Hydrelaps, Ephalophis, and Parahydrophis) form the sister group to the Hydrophis clade and have small paddles with poorly developed dorsal and ventral supports, consistent with their amphibious lifestyle. Overall, our results suggest that not only are the viviparous hydrophiines the only lineage of marine snakes to have acquired extremely large, skeletally supported caudal paddles but also that this innovation has occurred twice in the group in the past ~2-6 million years.     

Wake Vortex Structure Characteristics of a Flexible Oscillating Fin

We compute the wake of a two-dimensional and three-dimensional flexible fin in an unsteady flow field with heaving and pitching motions using FLUENT. Deflexion mode is used for a non-uniform cantilever beam with non-uniformly distributed load. The effect of chordwise deflexion length on the characteristics of propulsion is discussed for two-dimensional flexible fin. The thrust coefficient decreases, propulsive efficiency increases and the intensity of turbulence attenuates gradually as the deflexion length increases. For a three-dimensional flexible fin, the intensity of the vortex in the plane of symmetry is higher than that in the plane at 3/4 span length of the caudal fin. But the propulsive perform.ance achieved is not what we expected with the given deflexion mode.     

A comparison of pectoral fin ray morphology and its impact on fin ray flexural stiffness in labriform swimmers

The organization of tissues in appendages often affects their mechanical properties and function. In the fish family Labridae, swimming behavior is associated with pectoral fin flexural stiffness and morphology, where fins range on a continuum from stiff to relatively flexible fins. Across this diversity, pectoral fin flexural stiffness decreases exponentially along the length of any given fin ray, and ray stiffness decreases along the chord of the fin from the leading to trailing edge. In this study, we examine the morphological properties of fin rays, including the effective modulus in bending (E), second moment of area (I), segmentation, and branching patterns, and their impact on fin ray stiffness. We quantify intrinsic pectoral fin ray stiffness in similarly sized fins of two closely related species that employ fins of divergent mechanics, the flapping Gomphosus varius and the rowing Halichoeres bivittatus. While segmentation patterns and E were similar between species, measurements of I and the number of fin ray branch nodes were greater in G. varius than in H. bivittatus. A multiple regression model found that of these variables, I was always significantly correlated with fin ray flexural stiffness and that variation in I always explained the majority of the variation in flexural stiffness. Thus, while most of the morphological variables quantified in this study correlate with fin ray flexural stiffness, second moment of area is the greatest factor contributing to variation in flexural stiffness. Further, interspecific variation in fin ray branching pattern could be used as a means of tuning the effective stiffness of the fin webbing to differences in swimming behavior and hydrodynamics. The comparison of these results to other systems begins to unveil fundamental morphological features of biological beams and yields insight into the role of mechanical properties in fin deformation for aquatic locomotion.     

Morphology of the core fibrous layer of the cetacean tail fluke

The cetacean tail fluke blades are not supported by any vertebral elements. Instead, the majority of the blades are composed of a densely packed collagenous fiber matrix known as the core layer. Fluke blades from six species of odontocete cetaceans were examined to compare the morphology and orientation of fibers at different locations along the spanwise and chordwise fluke blade axes. The general fiber morphology was consistent with a three‐dimensional structure comprised of two‐dimensional sheets of fibers aligned tightly in a laminated configuration along the spanwise axis. The laminated configuration of the fluke blades helps to maintain spanwise rigidity while allowing partial flexibility during swimming. When viewing the chordwise sectional face at the leading edge and mid‐chord regions, fibers displayed a crossing pattern. This configuration relates to bending and structural support of the fluke blade. The trailing edge core was found to have parallel fibers arranged more dorso‐ventrally. The fiber morphology of the fluke blades was dorso‐ventrally symmetrical and similar in all species except the pygmy sperm whale (Kogia breviceps), which was found to have additional core layer fiber bundles running along the span of the fluke blade. These additional fibers may increase stiffness of the structure by resisting tension along their long spanwise axis.     

The functional anatomy of the shoulder of the savannah monitor lizard (Varanus exanthematicus)

The excursions of the scapulocoracoid and forelimb and the activity of 18 shoulder muscles were studied by simultaneous cineradiography and electromyography in Savannah Monitor lizards (Varanus exanthematicus) walking on a treadmill at speeds of 0.7–1.1 km/hour. During the propulsive phase, the humerus moves anteroposteriorly 40–55° and rotates a total of 30–40°. Simultaneously, the coracoid translates posteriorly along the tongue-and-groove coracosternal joint by a distance equivalent to about 40% the length of the coracoid. Biceps brachii, coraco-brachialis brevis and longus, the middle and posterior parts of the latissimus dorsi and pectoralis, serratus anterior, serratus anterior superficialis, subscapularis, supracoracoideus, and triceps usually become active during the late swing phase and continue activity throughout most or all of propulsion. The anterior part of the latissimus dorsi is active during the transition from propulsive to swing phases. Brachialis, deltoideus scapularis, levator scapulae, the anterior part of pectoralis, scapulo-humeralis posterior, and subcoracoideus are active primarily during the swing phase; they are occasionally active during propulsion. Deltoideus clavicularis, scapulo-humeralis posterior, sternocoracoideus, and the posterior part of the trapezius are biphasic, with activity in both the propulsive and swing phases. A number of shoulder muscles in Varanus exanthematicus and Didelphis virginiana (the Virginia opossum) are similar in attachments, in activity patterns with respect to phases of the step cycle, and in apparent actions. These similarities are interpreted as a pattern inherited from the ancestors of higher tetrapods. The sliding coracosternal joint permits an increase in step length without demanding greater excursion at the shoulder and elbow joints.     

Recombination rate and protein evolution in yeast

Background

Tetrapods exhibit great diversity in limb structures among species and also between forelimbs and hindlimbs within species, diversity which frequently correlates with locomotor modes and life history. We aim to examine the potential relation of changes in developmental timing (heterochrony) to the origin of limb morphological diversity in an explicit comparative and quantitative framework. In particular, we studied the relative time sequence of development of the forelimbs versus the hindlimbs in 138 embryos of 14 tetrapod species spanning a diverse taxonomic, ecomorphological and life-history breadth. Whole-mounts and histological sections were used to code the appearance of 10 developmental events comprising landmarks of development from the early bud stage to late chondrogenesis in the forelimb and the corresponding serial homologues in the hindlimb.

Results

An overall pattern of change across tetrapods can be discerned and appears to be relatively clade-specific. In the primitive condition, as seen in Chondrichthyes and Osteichthyes, the forelimb/pectoral fin develops earlier than the hindlimb/pelvic fin. This pattern is either retained or re-evolved in eulipotyphlan insectivores (= shrews, moles, hedgehogs, and solenodons) and taken to its extreme in marsupials. Although exceptions are known, the two anurans we examined reversed the pattern and displayed a significant advance in hindlimb development. All other species examined, including a bat with its greatly enlarged forelimbs modified as wings in the adult, showed near synchrony in the development of the fore and hindlimbs.

Conclusion

Major heterochronic changes in early limb development and chondrogenesis were absent within major clades except Lissamphibia, and their presence across vertebrate phylogeny are not easily correlated with adaptive phenomena related to morphological differences in the adult fore- and hindlimbs. The apparently conservative nature of this trait means that changes in chondrogenetic patterns may serve as useful phylogenetic characters at higher taxonomic levels in tetrapods. Our results highlight the more important role generally played by allometric heterochrony in this instance to shape adult morphology.