Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

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.     

Dynamic Modeling and Experiment of a Fish Robot with a Flexible Tail Fin

This paper presents the dynamic modeling of a flexible tail for a robotic fish. For this purpose firstly, the flexible tail was simplified as a slewing beam actuated by a driving moment. The governing equation of the flexible tail was derived by using the Euler-Bernoulli theory. In this equation, the resistive forces were estimated as a term analogous to viscous damping. Then, the modal analysis method was applied in order to derive an analytical solution of the governing equation, by which the relationship between the driving moment and the lateral movement of the flexible tail was described. Finally, simulations and experiments were carried out and the results were compared to verify the accuracy of the dynamic model. It was proved that the dynamic model of a fish robot with a flexible tail fin well explains the real behavior of robotic fish in underwater environment.     

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.     

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.     

Comparative morphology of the pectoral fin spine of the Persian sturgeon Acipenser persicus,the Russian sturgeon Acipenser gueldenstaedtii,and the Starry sturgeon Acipenser stellatus in Iranian waters of the Caspian Sea

The morphological characteristics of the pectoral fin spine were compared in three species of sturgeon, the Persian sturgeon (Acipenser persicus), the Russian sturgeon (Acipenser gueldenstaedtii), and the Starry sturgeon (Acipenser stellatus), all sampled from the Caspian Sea. On the basis of morphological characters of the pectoral fin spine, 62.2% of the individuals were correctly classified into separate groups. The cluster analysis also divided the three species into two major subgroups. Acipenser persicus and A. gueldenstaedtii were grouped together, suggesting a similar evolutionary basis. Significant morphological heterogeneity in pectoral fin spine characteristics was observed among the three sturgeon species. Principal component analysis identified the largest differences were in the pectoral fin spine size and the angle between distal pectoral fin spine and the horizontal line (A°). The first and second principal components (PC1 and PC2) of all observations accounted for 64.19% and 14.33% of the total variation, respectively. The combination of all analyses showed the relevance of applying pectoral fin spine shape for interspecific distinction of the three species of sturgeons.     

Pectoral Fin of the Megamouth Shark: Skeletal and Muscular Systems,Skin Histology,and Functional Morphology

This is the first known report on the skeletal and muscular systems, and the skin histology, of the pectoral fin of the rare planktivorous megamouth shark Megachasma pelagios. The pectoral fin is characterized by three features: 1) a large number of segments in the radial cartilages; 2) highly elastic pectoral fin skin; and 3) a vertically-rotated hinge joint at the pectoral fin base. These features suggest that the pectoral fin of the megamouth shark is remarkably flexible and mobile, and that this flexibility and mobility enhance dynamic lift control, thus allowing for stable swimming at slow speeds. The flexibility and mobility of the megamouth shark pectoral fin contrasts with that of fast-swimming sharks, such as Isurus oxyrhinchus and Lamna ditropis, in which the pectoral fin is stiff and relatively immobile.     

Gait Study and Pattern Generation of a Starfish-Like Soft Robot with Flexible Rays Actuated by SMAs

This paper presents the design and development of a starfish-like soft robot with flexible rays and the implementation of multi-gait locomotion using Shape Memory Alloy （SMA） actuators. The design principle was inspired by the starfish, which possesses a remarkable symmetrical structure and soft internal skeleton. A soft robot body was constructed by using 3D printing technology. A kinematic model of the SMA spring was built and developed for motion control according to displacement and force requirements. The locomotion inspired from starfish was applied to the implementation of the multi-ray robot through the flexible actuation induced multi-gait movements in various environments. By virtue of the proposed ray control patterns in gait transition, the soft robot was able to cross over an obstacle approximately twice of its body height. Results also showed that the speed of the soft robot was 6.5 times faster on sand than on a clammy rough terrain. These experiments demonstrated that the bionic soft robot with flexible rays actuated by SMAs and multi-gait locomotion in proposed patterns can perform successfully and smoothly in various terrains.     

Fibin, a novel secreted lateral plate mesoderm signal, is essential for pectoral fin bud initiation in zebrafish

We identified a novel secreted protein, fibin, in zebrafish, mice and humans. We inhibited its function in zebrafish embryos by injecting antisense fibin morpholino oligonucleotides. A knockdown of fibin function in zebrafish resulted in no pectoral fin bud initiation and abolished the expression of tbx5, which is involved in the specification of pectoral fin identification. The lack of pectoral fins in fibin-knockdown embryos was partially rescued by injection of fibin RNA. fibin was expressed in the lateral plate mesoderm of the presumptive pectoral fin bud regions. Its expression region was adjacent to that of tbx5. fibin expression temporally preceded tbx5 expression in presumptive pectoral fin bud regions, and not abolished in tbx5-knockdown presumptive fin bud regions. In contrast, fibin expression was abolished in retinoic acid signaling-inhibited or wnt2b-knockdown presumptive fin bud regions. These results indicate that fibin is a secreted signal essential for pectoral fin bud initiation in that it potentially acts downstream of retinoic acid and wnt signaling and is essential for tbx5 expression. The present findings have revealed a novel secreted lateral plate mesoderm signal essential for fin initiation in the lateral plate mesoderm.     

Re‐evaluation of batoid pectoral morphology reveals novel patterns of diversity among major lineages

Batoids (Chondrichthyes: Batoidea) are a diverse group of cartilaginous fishes which comprise a monophyletic sister lineage to all neoselachians or modern sharks. All species in this group possess anteroposteriorly expanded‐pectoral fins, giving them a unique disc‐like body form. Reliance on pectoral fins for propulsion ranges from minimal (sawfish) to almost complete dependence (skates and rays). A recent study on the diversity of planform pectoral fin shape in batoids compared overall patterns of morphological variation within the group. However, inconsistent pectoral homology prevented the study from accurately representing relationships within and among major batoid taxa. With previous work in mind, we undertook an independent investigation of pectoral form in batoids and evaluated the implications of shape diversity on locomotion and lifestyle, particularly in the skates (Rajoidei) and rays (Myliobatoidei). We used geometric morphometrics with sliding semilandmarks to analyze pectoral fin outlines and also calculate fin aspect ratios (AR), a functional trait linked to locomotion. In agreement with previous work, our results indicated that much of the evolution of batoid pectoral shape has occurred along a morphological axis that is closely related to AR. For species where kinematic data were available, both shape and AR were associated with swimming mode. This work further revealed novel patterns of shape variation among batoids, including strong bimodality of shape in rays, an intermediate location of skate species in the morphospace between benthic/demersal and pelagic rays, and approximately parallel shape trajectories in the benthic/demersal rays and skates. Finally, manipulation of landmarks verified the need for a consistent and accurate definition of homology for the outcome and efficacy of analyses of pectoral form and function in batoids. J. Morphol. 277:482–493, 2016. © 2016 Wiley Periodicals, Inc.     

Micro-anatomy of the pectoral fin in blennies (Blenniini, Blennioidea, Teleostei)

Blennioid fishes show a highly differentiated pectoral fin, which they use to cling to the substrate. The lower part of the pectoralis, comprising about four to six fin rays, forms a hook-field with specific anatomical features: (1) the rim of the fin web has a saw-like appearance, because it extends from the tip of a fin ray to the shaft ofthe upper of two neighbouring fin rays, (2) the outer half of the bony fin ray carries a lepidotrichal cord composed of fibrocytes, collagen, elastic fibres and acidic GAGS, (3) the epidermis overlying the lepidotrichal cord is differentiated in terms of cyto-architecture and forms a conspicuous cuticle. The upper part of the pectoral fin does not show any obvious specializations and is used for swimming and undulation. The vascularization of the fin originates from a stem vessel which gives rise to five branches, each supplying two or three neighbouring fin rays. Each fin ray is accompanied by a single arterial vessel at its upper edge. No vessels are found in the space between the bony fin ray halves. The morphology of the shoulder girdle and pectoral fin shows only little variation among the four species of Blenniini studied. Most remarkable is the fusion of the coracoid with the cleithrum, loss of one element of the suspensorium and the absence of branched fin rays. The possible relevance of the Blennioid pectoral fin as a model for the origin of morphological novelties in connection with functional specializations is discussed.     

Diversity of pectoral fin structure and function in fishes with labriform propulsion

Aquatic propulsion generated by the pectoral fins occurs in many groups of perciform fishes, including numerous coral reef families. This study presents a detailed survey of pectoral fin musculoskeletal structure in fishes that use labriform propulsion as the primary mode of swimming over a wide range of speeds. Pectoral fin morphological diversity was surveyed in 12 species that are primarily pectoral swimmers, including members of all labroid families (Labridae, Scaridae, Cichlidae, Pomacentridae, and Embiotocidae) and five additional coral reef fish families. The anatomy of the pectoral fin musculature is described, including muscle origins, insertions, tendons, and muscle masses. Skeletal structures are also described, including fin shape, fin ray morphology, and the structure of the radials and pectoral girdle. Three novel muscle subdivisions, including subdivisions of the abductor superficialis, abductor profundus, and adductor medialis were discovered and are described here. Specific functional roles in fin control are proposed for each of the novel muscle subdivisions. Pectoral muscle masses show broad variation among species, particularly in the adductor profundus, abductor profundus, arrector dorsalis, and abductor superficialis. A previously undescribed system of intraradial ligaments was also discovered in all taxa studied. The morphology of these ligaments and functional ramifications of variation in this connective tissue system are described. Musculoskeletal patterns are interpreted in light of recent analyses of fin behavior and motor control during labriform swimming. Labriform propulsion has apparently evolved independently multiple times in coral reef fishes, providing an excellent system in which to study the evolution of pectoral fin propulsion.     

Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic vs. pelagic station-holding

Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column.     

The dorsal fin engine of the seahorse (Hippocampus sp.)

The muscles, fin ray joints, and supporting structures underlying the dorsal fin are described for two seahorse species: Hippocampus zosterae and Hippocampus erectus. A fan-shaped array of cartilaginous bones, the pterigiophores, form the internal supporting structure of the dorsal fin. Each pterigiophore is composed of a proximal radial that extends from a vertebra to the dorsal side of the animal, where it fuses to a middle radial. The middle radials fuse with each other to form a dorsal ridge upon which sit the spheroidal distal radials. Each distal radial articulates with a fin ray on its dorsal side and is attached to the dorsal ridge on its ventral side by a material that has been histologically identified as elastic cartilage. Together these connections form a two-axis joint that permits elevation, depression, and inclination of the ray. Each fin ray is actuated by two bilateral pairs of muscles, an anterior pair of inclinators, and a posterior pair of depressors. The anteriormost fin ray is actuated by three bilateral pair of muscles, the inclinators, the depressors, and a pair of elevator muscles that are positioned anterior to the inclinators. Preliminary examinations of the ray joints of the pectoral and anal fins of adult H. zostera and the pectoral fins of newborn H. erectus revealed structures similar to that seen in the dorsal fins. To further explore the structure and function of the dorsal fin gross dissections and simple functional tests were performed on H. erectus and H. barbouri and behavioral observations were made of all three species plus Hippocampus kuda.     

棒花鱼形态特征的两性异形和雌性个体生育力

测定了棒花鱼(Abbottina rivularis)繁殖期形态特征包括体长、头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、胸鳍腹鳍间距、尾柄长、尾鳍长和体重的两性异形和雌性个体生育力。结果表明,雄性个体的数量显著多于雌性个体,雄性个体的体长显著大于雌性个体。特定体长的雌性个体的胸鳍腹鳍间距显著大于雄性个体,头长、头宽、头高、眼间距、鼻间距、背鳍基长、胸鳍长、尾柄长和尾鳍长显著小于雄性个体,雌雄两性体重不存在显著差异。棒花鱼的怀卵数量与体长和体重回归关系显著。偏相关分析显示,当控制第三者恒定时,怀卵数量与体长和体重呈正相关但不显著。棒花鱼存在个体大小和其他局部特征显著的两性异形,雌性个体主要通过腹腔容积的增加提高个体生育力。棒花鱼形态特征的两性异形是性选择和生育力选择共同作用的结果。     

新型有原始肢突胴甲鱼的发现及胴甲鱼早期演化的初步探讨

本文记述的曲靖始突鱼(Procondylolepis qujingensis gen.et sp.nov.)是近几年在云南曲靖早泥盆世地层中发现的有肢突胴甲鱼一原始类型。它和已知胴甲鱼(包括早泥盆世无肢突的和中、晚泥盆世有“盔”状肢突的)不同的最大特点是在其肩带与胸鳍相接的肩关节处有原始的肢突和简单的关节窝;胸鳍甲近端的关节区很小。它展现出胴甲鱼类这一高度特化、长期使人迷惑不解的肢突,在胴甲鱼演化史上发展变化的梗概,填补了肢突从无到有中间的缺环,使人了解到胸鳍的具体结构。文中主要根据肢突的有无和特化程度等,对胴甲鱼早期演化史作了初步探讨,将胴甲鱼分为无肢突超目(Abrachicondylia)和有肢突超目(Brachicondylia)两大部分。始突鱼则代表有肢突超目一早期成员。     

Development of the paired fins in the paddlefish, Polyodon spathula

In Polyodon spathula, the pectoral fin radials, with the exception of the metapterygium, are derived from the decomposition of a single continuous cartilage fin plate that is continuous with the scapulocoracoid. This cartilage sheet develops two interior splits to form three precursor pieces, and these decompose in a predictable way to generate the propterygium and radials. The metapterygium is an extension of the scapulocoracoid that segments off of it during early development. To our knowledge, this has not been reported for acipenserids or other basal actinopterygians. In teleosts, the proximal radials also develop from the "break up" of an initially continuous paddle-like sheet of cartilage along the posterior edge of the scapulocoracoid, and in Polypterus and sharks a similar pattern holds. Thus, the pattern observed in Polyodon may represent the basal developmental condition for the gnathostome pectoral fin. The process underlying development of the superficially similar cartilages of the pelvic and pectoral fins is different. In the pectoral fin, the metapterygium is segmented off of the scapulocoracoid and other radials form from the decomposition of the cartilage plate. In contrast, individual rod-like basipterygial elements form in a close one-to-one correspondence with the middle radials of the pelvic fin, but later fuse to form an anterior element that is branched in appearance. To evaluate further claims of similarity among the pectoral and pelvic fin elements of various fishes, the course of the development of these structures must be observed. The pectoral fin and girdle in Polyodon ossifies in a different sequence than that proposed as ancestral (and highly conserved) for actinopterygians: the supracleithrum ossifies significantly before the cleithrum. The later ossification of the cleithrum in Polyodon may be related to the primary use of the caudal fin vs. the pectoral fins in their locomotion.     

西藏双须叶须鱼早期发育特征

通过双须叶须鱼(Ptychobarbus dipogon Regan)早期发育特征研究, 旨在为该鱼的科学保护和合理开发提供技术支撑。结果显示: 双须叶须鱼卵径3.7—3.9 mm, 吸水后的卵径可达5.1—5.3 mm。在水温10℃左右的条件下, 经历336.02h孵化出膜。根据胚胎的外部形态特征可将胚胎发育分为准备卵裂阶段、卵裂阶段、囊胚阶段、原肠阶段、神经胚阶段、器官分化阶段、孵化阶段共7个阶段34个时期。初孵仔鱼全长12.4 mm, 第1天体色素出现, 胸鳍上翘, 鳃盖骨出现, 下颌原基出现; 第2天鳃弓原基出现; 第3天消化道出现, 肝胰脏原基出现; 第4天鳃耙出现, 体表色素细胞带出现; 第5天口凹形成, 鳃丝形成; 第6天胸鳍褶, 背鳍褶, 腹鳍褶出现; 第7天鼻凹出现, 星芒状色素团出现; 第9天鳔前原基出现; 第11天尾鳍鳍条开始出现, 胸鳍开始颤动; 第13天鳔1室出现, 半规管形成; 第17天背鳍分化出来; 第21天腹部鳍褶变大, 舌颌骨清晰可见; 第28天脾脏出现; 第33天出现腹鳍鳍条; 第34天鳞片出现; 第85天稚鱼的形态与成鱼无异。双须叶须鱼是已报道裂腹鱼类卵径最大, 较四大家鱼卵周隙小, 是对高原隆起所导致的高寒自然环境的一种适应。     

A new deep‐water chimaerid species, Hydrolagus lusitanicus n. sp., from off mainland Portugal with a proposal of a new identification key for the genus Hydrolagus(Holocephali: Chimaeridae) in the north‐east Atlantic

A new chimaerid species closely related to Hydrolagus pallidus is described from 13 specimens captured on the Portuguese continental slope (north‐east Atlantic) by commercial longliners at depths of c . 1600 m. The new species is large‐bodied with a rose to light brown body colouration. Hydrolagus lusitanicus n.sp. presents a combination of external morphological characters that allow it to be clearly differentiated from its congeners, in particular, the ratios of pectoral fin length: pectoral fin width and pelvic fin length: pelvic fin width; in having a serrated posterior edge of the dorsal spine; a high number of ridges in the dental plates. Additionally in males, there are differences on the shape and number of hooks in the frontal tenacula, on the number of spines in the prepelvic tenacula and on the length of the pectoral fin margin.