Use of biorobotic models of highly deformable fins for studying the mechanics and control of fin forces in fishes

Bony fish swim with a level of agility that is unmatched in human-developed systems. This is due, in part, to the ability of the fish to carefully control hydrodynamic forces through the active modulation of the fins' kinematics and mechanical properties. To better understand how fish produce and control forces, biorobotic models of the bluegill sunfish's (Lepomis macrochirus) caudal fin and pectoral fins were developed. The designs of these systems were based on detailed analyses of the anatomy, kinematics, and hydrodynamics of the biological fins. The fin models have been used to investigate how fin kinematics and the mechanical properties of the fin-rays influence propulsive forces and to explore kinematic patterns that were inspired by biological motions but that were not explicitly performed by the fish. Results from studies conducted with the fin models indicate that subtle changes to the kinematics and mechanical properties of fin rays can significantly impact the magnitude, direction, and time course of the 3D forces used for propulsion and maneuvers. The magnitude of the force tends to scale with the fin's stiffness, but the direction of the force is not invariant, and this causes disproportional changes in the magnitude of the thrust, lift, and lateral components of force. Results from these studies shed light on the multiple strategies that are available to the fish to modulate fin forces.     

Musculoskeletal morphology and regionalization within the dorsal and anal fins of bluegill sunfish (Lepomis macrochirus)

Ray‐finned fishes actively control the shape and orientation of their fins to either generate or resist hydrodynamic forces. Because of the emergent mechanical properties of their segmented, bilaminar fin rays (lepidotrichia), and actuation by multiple muscles, fish can control the rigidity and curvature of individual rays independently, thereby varying the resultant forces across the fin surfaces. Expecting that differences in fin‐ray morphology should reflect variation in their mechanical properties, we measured several musculoskeletal features of individual spines and rays of the dorsal and anal fins of bluegill sunfish, Lepomis macrochirus, and assessed their mobility and flexibility. We separated the fin‐rays into four groups based on the fin (dorsal or anal) or fin‐ray type (spine or ray) and measured the length of the spines/rays and the mass of the three median fin‐ray muscles: the inclinators, erectors and depressors. Within the two ray groups, we measured the portion of the rays that were segmented vs. unsegmented and branched vs. unbranched. For the majority of variables tested, we found that variations between fin‐rays within each group were significantly related to position within the fin and these patterns were conserved between the dorsal and anal rays. Based on positional variations in fin‐ray and muscle parameters, we suggest that anterior and posterior regions of each fin perform different functions when interacting with the surrounding fluid. Specifically, we suggest that the stiffer anterior rays of the soft dorsal and anal fins maintain stability and keep the flow across the fins steady. The posterior rays, which are more flexible with a greater range of motion, fine‐tune their stiffness and orientation, directing the resultant flow to generate lateral and some thrust forces, thus acting as an accessory caudal fin. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Experimental hydrodynamics of fish locomotion: functional insights from wake visualization

Despite enormous progress during the last twenty years in understandingthe mechanistic basis of aquatic animal propulsion—a taskinvolving the construction of a substantial data base on patternsof fin and body kinematics and locomotor muscle function—thereremains a key area in which biologists have little information:the relationship between propulsor activity and water movementin the wake. How is internal muscular force translated intoexternal force exerted on the water? What is the pattern offluid force production by different fish fins (e.g., pectoral,caudal, dorsal) and how does swimming force vary with speedand among species? These types of questions have received considerableattention in analyses of terrestrial locomotion where forceoutput by limbs can be measured directly with force plates.But how can forces exerted by animals moving through fluid bemeasured? The advent of digital particle image velocimetry (DPIV)has provided an experimental hydrodynamic approach for quantifyingthe locomotor forces of freely moving animals in fluids, andhas resulted in significant new insights into the mechanismsof fish propulsion. In this paper we present ten "lessons learned"from the application of DPIV to problems of fish locomotionover the last five years. (1) Three-dimensional DPIV analysisis critical for reconstructing wake geometry. (2) DPIV analysisreveals the orientation of locomotor reaction forces. (3) DPIVanalysis allows calculation of the magnitude of locomotor forces.(4) Swimming speed can have a major impact on wake structure.(5) DPIV can reveal interspecific differences in vortex wakemorphology. (6) DPIV analysis can provide new insights intothe limits to locomotor performance. (7) DPIV demonstrates thefunctional versatility of fish fins. (8) DPIV reveals hydrodynamicforce partitioning among fins. (9) DPIV shows that wake interactionamong fins may enhance thrust production. (10) Experimentalhydrodynamic analysis can provide insight into the functionalsignificance of evolutionary variation in fin design.     

Terrestrial locomotion in the climbing perch, Anabas testudineus (Bloch) (Anabantidea, Pisces)

Climbing perches use the tail for forward propulsion when on land. The spiny gill covers are used to obtain purchase on the substratum. Unusually amongst teleosts, the opercular and subopercular bone are not bound strongly together into a single operculum. Instead they are joined only by a thin, flexible membrane, so that the fish has two sections of the gill cover hinged separately; the opercular on the suspensorium and the subopercular on the rearmost part of the lower jaw. The gill covers open very widely and the subopercular rotates ventrally as well as laterally. Two modes of locomotion are employed. Usually Anabus adopts a near-upright posture and alternately drives the left and right spiny suboperculars into the substratum, using the tail to vault over the subopercular (which acts in the manner of a short vaulting pole). During vigorous movement the fish may leave the substratum altogether during the vault. Fatigued fish, or fish which have fallen during climbing, move on their sides, using the tail for propulsion and a single spiny opercular which is repeatedly driven into the substratum. Maximum speeds observed on land corresponded to 1.8 body lengths S^?1. No evidence of involvement of the pelvic fins in locomotion was obtained; the pectorals seemed only to help in tilting the head from side to side and did not contribute significantly to forward propulsion. On mud and tree bark the fish were laterally unstable, often resting on their sides. Grassy substrata provided support and permitted a more upright posture. Slope climbing ability was restricted to 25° when climbing tree bark and 30° when climbing grass. Coupled with lateral instability these observations disqualify the species from any tree-climbing capacity. Anabas can climb vertical obstacles of at least half body length in height by pushing the head against the obstacle, using the spiny gill covers for purchase until one or other subopercular hooks over the top of the barrier. The subopercular is then used as a fulcrum for an upwards vault to clear the obstacle.     

Pectoral fin morphology of batoid fishes (Chondrichthyes: Batoidea): Explaining phylogenetic variation with geometric morphometrics

The diverse cartilaginous fish lineage, Batoidea (rays, skates, and allies), sister taxon to sharks, comprises a huge range of morphological diversity which to date remains unquantified and unexplained in terms of evolution or locomotor style. A recent molecular phylogeny has enabled us to confidently assess broadscale aspects of morphology across Batoidea. Geometric morphometrics quantifies the major aspects of shape variation, focusing on the enlarged pectoral fins which characterize batoids, to explore relationships between ancestry, locomotion and habitat. A database of 253 specimens, encompassing 60 of the 72 batoid genera, reveals that the majority of morphological variation across Batoidea is attributable to fin aspect‐ratio and the chordwise location of fin apexes. Both aspect‐ratio and apex location exhibit significant phylogenetic signal. Standardized independent linear contrast analysis reveals that fin aspect‐ratio can predict locomotor style. This study provides the first evidence that low aspect‐ratio fins are correlated with undulatory‐style locomotion in batoids, whereas high aspect‐ratio fins are correlated with oscillatory locomotion. We also show that it is phylogeny that determines locomotor style. In addition, body‐ and caudal fin‐locomotors are shown to exhibit low aspect‐ratio fins, whereas a pelagic lifestyle correlates with high aspect‐ratio fins. These results emphasize the importance of phylogeny in determining batoid pectoral fin shape, however, interactions with other constraints, most notably locomotor style, are also highlighted as significant. J. Morphol. 275:1173–1186, 2014. © 2014 Wiley Periodicals, Inc.     

Functional morphology of the fin rays of teleost fishes

Ray‐finned fishes are notable for having flexible fins that allow for the control of fluid forces. A number of studies have addressed the muscular control, kinematics, and hydrodynamics of flexible fins, but little work has investigated just how flexible ray‐finned fish fin rays are, and how flexibility affects their response to environmental perturbations. Analysis of pectoral fin rays of bluegill sunfish showed that the more proximal portion of the fin ray is unsegmented while the distal 60% of the fin ray is segmented. We examined the range of motion and curvatures of the pectoral fin rays of bluegill sunfish during steady swimming, turning maneuvers, and hovering behaviors and during a vortex perturbation impacting the fin during the fin beat. Under normal swimming conditions, curvatures did not exceed 0.029 mm^?1 in the proximal, unsegmented portion of the fin ray and 0.065 mm^?1 in the distal, segmented portion of the fin ray. When perturbed by a vortex jet traveling at approximately 1 ms^?1 (67 ± 2.3 mN s.e. of force at impact), the fin ray underwent a maximum curvature of 9.38 mm^?1. Buckling of the fin ray was constrained to the area of impact and did not disrupt the motion of the pectoral fin during swimming. Flexural stiffness of the fin ray was calculated to be 565 × 10^?6 Nm². In computational fluid dynamic simulations of the fin‐vortex interaction, very flexible fin rays showed a combination of attraction and repulsion to impacting vortex dipoles. Due to their small bending rigidity (or flexural stiffness), impacting vortices transferred little force to the fin ray. Conversely, stiffer fin rays experienced rapid small‐amplitude oscillations from vortex impacts, with large impact forces all along the length of the fin ray. Segmentation is a key design feature of ray‐finned fish fin rays, and may serve as a means of making a flexible fin ray out of a rigid material (bone). This flexibility may offer intrinsic damping of environmental fluid perturbations encountered by swimming fish. J. Morphol. 274:1044–1059, 2013. © 2013 Wiley Periodicals, Inc.     

A Three-Dimensional Kinematics Analysis of a Koi Carp Pectoral Fin by Digital Image Processing

Pectoral fins fascinate researchers for their important role in fish maneuvers. By possessing a complicated flexible structure with several fin rays made by a thin film, the fin exhibits a three-dimensional (3D) motion. The complex 3D fin kinematics makes it challenging to study the performance of pectoral fin. Nevertheless, a detailed study on the 3D motion pattern of pectoral fins is necessary to the design and control of a bio-inspired fin rays. Therefore, a highspeed photography system is introduced in this paper to study the 3D motion of a Koi Carp by analyzing the two views of its pectoral fin simultaneously. The key motions of the pectoral fins are first captured in both hovering and retreating. Next, the 3D configuration of the pectoral fins is reconstructed by digital image processing, in which the movement of fin rays during fish retreating and hovering is obtained. Furthermore, the method of Singular Value Decomposition (SVD) is adopted to extract the basic motion patterns of pectoral fins from extensive image sequences, i.e. expansion, bending, cupping, and undulation. It is believed that the movement of the fin rays and the basic patterns of the pectoral fins obtained in the present work can provide a good foundation for the development and control of bionic flexible pectoral fins for underwater propeller.     

Numerical and Experimental Research on Modular Oscillating Fin

Fishes are famous for their ability to position themselves accurately even in turbulent flows. This ability is the result of the coordinated movement of fins which extend from the body. We have embarked on a research program designed to develop an agile and high efficient biologically inspired robotic fish based on the performance of hybrid mechanical fms. To accomplish this goal, a mechanical ray-like fin actuated by Shape Memory Alloy （SMA） is developed, which can realize both oscillatory locomotion and undulatory locomotion. We first give a brief introduction on the mechanical structure of our fin and then carry out theoretic analysis on force generation. Detailed information of these theoretical results is later revealed by Computational Huid Dynamic （CFD）, and is final validated by experiments. This robotic fin has potential application as a propulsor for future underwater vehicles in addition to being a valuable scientific instrument.     

Morphological aspects of the development of swimming and feeding functions in the milkfishChanos chanos

Development of swimming and feeding abilities based on morphological development of larval and early juvenileChanos chanos was investigated. In larvae smaller than about 6.5 mm SL, mechanical supports of fins and branchial arches were in a primordial stage of development. Supports and rays of the vertical fins and branchial arches rapidly developed from 6.5 mm SL, and all components appeared by about 10.5 mm SL. Thereafter body depth proportion changed and the supports and rays of the paired fins and gill-rakers developed. These developmental events were nearly or totally completed by about 17 mm SL, and we concluded that the larvae transformed to juveniles at this size. By this time, the mode of swimming of the fish shifted from undulating locomotion to caudal propulsion and that of feeding from swallowing paniculate food to filtering and concentrating substrate food matters using gill-rakers and the epibranchial organ. One of the most characteristic, and well-known, phenomena in the life history ofChanos chanos is the mass occurrence in the surf zone of postlarvae of a limited size range. In view of the scheme of the development of mechanical supports of the body and fins, they may acquire a swimming ability strong enough to move against the current only upon reaching about 10.5 mm SL, and if active shoreward migration of the larvae occurs, it is only during the late period of their journey from the spawning grounds to the shore. The sudden disappearance from the surf zone of larvae larger than 15–16 mm SL is obviously related to a change in food habit.     

Effect of Flow Turbulence on Swimming Speed of Fish

The effect of flow turbulence on the swimming speed was studied in perch (Perca fluviatilis) with different body length. The critical flow rate was used as an index of fish swimming performance. The longer was the fish, the higher turbulence was required to decrease the critical flow rate. The mechanism of turbulence impact on fish locomotion relied on the vortex structure of the flow. The torque produced by hydrodynamic forces in a vortex favors fish overturn and loss of balance. Such effect of turbulence was observed when the sizes of the vortex and fish body were similar. The fish uses the pectoral fins to restore the balance, which increases their hydraulic resistance and, together with energy expenditure for spatial balance control, decreases the swimming speed.     

Normal growth of the "see-through" medaka

The see-through stock in the medaka Oryzias latipes, causes pigments to be absent from the whole body and has a transparent body in the adult stage as well as during embryonic stages. To establish a standard table of growth stages for this model fish, morphological features were examined during the growing period from hatching to adulthood. The main observations were performed on morphological changes in external and internal organs that could be seen through the body wall of the living fish during growth. Finally, five growth stages from just after hatching to the adult stage were defined on the basis of synchronized or definite changes in morphology as follows: (1) stage 40 in which the nodes (joints) in bony rays of the caudal and pectoral fins first appear, (2) the stage 41 in which the ribs and the anal, dorsal and ventral fins are formed by degeneration of the membrane fin folds, as recognized by the first appearance of nodes in the fin rays of the anal, pectoral and dorsal fins, and the parallel distribution of the dorsal artery and ventral vein of the tail, (3) stage 42 in which the 2-spiral pattern of the gut, the ray nodes in the ventral fins, and the scales first appear, (4) stage 43 in which early secondary sexual characters such as urinogenital protruberances (female) and papillar processes (male) appear, (5) stage 44 in which the 3-spiral pattern of the gut and the papillar process on the 2nd ray of pectral fins (male) appear.     

Locomotion and adhesion of polymorphonuclear leukocytes effects of the supporting substratum

Contact angle measurements have been used to correlate surface hydrophobicity of a supporting substratum with adhesion and locomotion of polymorphonuclear leukocytes. The binding of human serum albumin, a well-known chemokinetic substance, to hydrophilic glass slides gave rise to hydrophobic surfaces with adhesive properties conducive, to cell polarization thus allowing cell locomotion. Parallel contact angle and cell adhesion measurements suggested that albumin modified the cellsubstratum interaction by increasing the van der Waals forces of attraction and reducing the electrostatic forces. By allowing cells to adhere to a hydrophobic surface (siliconized glass), it was found that protein could be omitted from in vitro test systems for leukocyte locomotion. It is suggested that quantitatively equal cell adhesion values may, depending on the type of attraction forces working in adhesion to the substratum, result in different locomotion patterns.     

Locomotion and adhesion of polymorphonuclear leukocytes. Effects of the supporting substratum

Contact angle measurements have been used to correlate surface hydrophobicity of a supporting substratum with adhesion and locomotion of polymorphonuclear leukocytes. The binding of human serum albumin, a well-known chemokinetic substance, to hydrophilic glass slides gave rise to hydrophobic surfaces with adhesive properties conductive to cell polarization, thus allowing cell locomotion. Parallel contact angle and cell adhesion measurements suggested that albumin modified the cell-substratum interaction by increasing the van der Waals forces of attraction and reducing the electrostatic forces. By allowing cells to adhere to a hydrophobic surface (siliconized glass), it was found that protein could be omitted from in vitro test systems for leukocyte locomotion. It is suggested that quantitatively equal cell adhesion values may, depending on the type of attraction forces working in adhesion to the substratum, result in different locomotion patterns.     

Vortex interactions with flapping wings and fins can be unpredictable

As they fly or swim, many animals generate a wake of vortices with their flapping fins and wings that reveals the dynamics of their locomotion. Previous studies have shown that the dynamic interaction of vortices in the wake with fins and wings can increase propulsive force. Here, we explore whether the dynamics of the vortex interactions could affect the predictability of propulsive forces. We studied the dynamics of the interactions between a symmetrically and periodically pitching and heaving foil and the vortices in its wake, in a soap-film tunnel. The phase-locked movie sequences reveal that abundant chaotic vortex-wake interactions occur at high Strouhal numbers. These high numbers are representative for the fins and wings of near-hovering animals. The chaotic wake limits the forecast horizon of the corresponding force and moment integrals. By contrast, we find periodic vortex wakes with an unlimited forecast horizon for the lower Strouhal numbers (0.2–0.4) at which many animals cruise. These findings suggest that swimming and flying animals could control the predictability of vortex-wake interactions, and the corresponding propulsive forces with their fins and wings.     

Biomechanics of swimming in the pufferfish Diodon holocanthus: propulsive momentum enhancement is an adaptation for thrust production in an undulatory median and paired-fin swimmer

A form of large-amplitude elongated-body theory appropriate for the analysis of undulatory fins attached to a rigid body of elliptical section suggests a benefit due to momentum enhancement relative to the fins on their own. This theoretical prediction is experimentally confirmed for the first time. Theoretical momentum enhancement factors for Diodon holocanthus (2.2 and 2.7 for the median and pectoral fins, respectively) compared well to inferred thrust values determined from particle-image velocimetry (PIV) wake measurements (2.2-2.4 and 2.7-2.9). Caudal fin mean theoretical thrust was not significantly different from measured (PIV) values (n = 24, P > 0.05), implying no momentum enhancement. Pectoral-fin thrust was half that of the median and caudal fins due to high fin-jet angles, low circulation and momentum. Average total fin thrust and fish drag were not significantly different (n = 24, P > 0.05). Vortex rings generated by the fins were elliptical, with size dependent on fin chord and stroke amplitude. Hydrodynamic advantages (thrust enhancement at no cost to hydrodynamic efficiency, reduction of side forces minimizing energy wasting yawing motions and body drag) are probably common among rigid-bodied organisms propelled by undulatory fins. A trade-off between momentum enhancement and the rate of momentum generation (thrust force) sets a practical limit to the former. For small fins whilst momentum enhancement is high, absolute thrust is low. In addition, previously suggested limitations on thrust enhancement set by reductions in propulsive force associated with progressive reductions in fin wavelength are found to be biologically unrealistic.     

Experimental Hydrodynamics and Evolution: Function of Median Fins in Ray-finned Fishes

The median fins of fishes consist of the dorsal, anal, and caudal fins and have long been thought to play an important role in generating locomotor force during both steady swimming and maneuvering. But the orientations and magnitudes of these forces, the mechanisms by which they are generated, and how fish modulate median fin forces have remained largely unknown until the recent advent of Digital Particle Image Velocimetry (DPIV) which allows empirical analysis of force magnitude and direction. Experimental hydrodynamic studies of median fin function in fishes are of special utility when conducted in a comparative phylogenetic context, and we have examined fin function in four ray-finned fish clades (sturgeon, trout, sunfish, and mackerel) with the goal of testing classical hypotheses of fin function and evolution. In this paper we summarize two recent technical developments in DPIV methodology, and discuss key recent findings relevant to median fin function. High-resolution DPIV using a recursive local-correlation algorithm allows quantification of small vortices, while stereo-DPIV permits simultaneous measurement of x, y, and z flow velocity components within a single planar light sheet. Analyses of median fin wakes reveal that lateral forces are high relative to thrust force, and that mechanical performance of median fins (i.e., thrust as a proportion of total force) averages 0.35, a surprisingly low value. Large lateral forces which could arise as an unavoidable consequence of thrust generation using an undulatory propulsor may also enhance stability and maneuverability. Analysis of hydrodynamic function of the soft dorsal fin in bluegill sunfish shows that a thrust wake is generated that accounts for 12% of total thrust and that the thrust generation by the caudal fin may be enhanced by interception of the dorsal fin wake. Integration of experimental studies of fin wakes, computational approaches, and mechanical models of fin function promise understanding of instantaneous forces on fish fins during the propulsive cycle as well as exploration of a broader locomotor design space and its hydrodynamic consequences.     

Does the common topknot Zeugopterus punctatus (Teleosteii: Pleuronectiformes: Scophthalmidae) use a novel Venturi-effect attachment mechanism? A testable hypothesis

Common topknots (Zeugopterus punctatus) attach to vertical rock surfaces and overhangs. It has been speculated that attachment is by a suction cup, with the median (anal, dorsal) fins providing a peripheral seal. Here the authors propose that the attachment is actually based on a Venturi effect. The rear portions of the median fins continually move in a fan-like fashion (at c. 4 cycles per second). This movement produces a tailward fluid flow that ventilates the shallow underbody space between the fish and its rocky substratum. The anterior portions of the median fins seal the space laterally, but the space is open anterior (beneath the raised head) and posterior to the sea. The mid-underbody space likely has a lower cross-sectional area than does the front intake or rear exit, so flow should be faster (and pressure lower) within it than outside, thus providing pressure gradient suction. Topknots attach to rough and heavily biofouled surfaces, presumably because the high numbers of fin rays and their associated membranes plus fine muscle control allow effective sealing. The attachment ability is shared by all members of the flatfish tribe Phrynorhombini; it can be related to anatomical peculiarities and constitutes a probable synapomorphy for this clade.     

The origins of adipose fins: an analysis of homoplasy and the serial homology of vertebrate appendages

Adipose fins are appendages found on the dorsal midline between the dorsal and caudal fins in more than 6000 living species of teleost fishes. It has been consistently argued that adipose fins evolved once and have been lost repeatedly across teleosts owing to limited function. Here, we demonstrate that adipose fins originated repeatedly by using phylogenetic and anatomical evidence. This suggests that adipose fins are adaptive, although their function remains undetermined. To test for generalities in the evolution of form in de novo vertebrate fins, we studied the skeletal anatomy of adipose fins across 620 species belonging to 186 genera and 55 families. Adipose fins have repeatedly evolved endoskeletal plates, anterior dermal spines and fin rays. The repeated evolution of fin rays in adipose fins suggests that these fins can evolve new tissue types and increased structural complexity by expressing fin-associated developmental modules in these new territories. Patterns of skeletal elaboration differ between the various occurrences of adipose fins and challenge prevailing hypotheses for vertebrate fin origin. Adipose fins represent a powerful and, thus far, barely studied model for exploring the evolution of vertebrate limbs and the roles of adaptation and generative biases in morphological evolution.     

Cutaneous taste buds in gadoid fishes

Cutaneous taste buds occurred on the head and fins in five species of juvenile gadoid fishes from the west of Scotland, but there were significant differences in their density between regions on the fish and between species for individual regions. The highest taste bud densities were recorded on the edge of the anterior naris flap, the barbel, pelvic fin rays, snout tip and upper lip. Cod Gadus morhua and poor cod Trisopterus minutus had significantly higher taste bud densities on the first two pelvic fin rays than the other species. This appears to correspond with their more benthic lifestyle, in which the pelvic fins are frequently trailed over the sea bed when searching for prey.