Caudal Fin Locomotion in Ray-finned Fishes: Historical and Functional Analyses

SYNOPSIS. Over the last 20 years, considerable progress hasbeen made in quantifying the movement of the body during locomotionby aquatic vertebrates, and in defining the role of axial musculaturein producing these kinematic patterns. Relatively little isknown, however, about how specific internal structural featuresof the axial system in fishes affect body kinematics, and howsuch structural and functional features have changed duringevolution. The major theme of this paper is that historical,phylogenetic patterns in the axial musculoskeletal system needto be integrated with experimental and functional data in orderto understand the design of the locomotor apparatus in vertebrates.To illustrate this proposition, the evolution of the tail inray-finned fishes is presented as a case study in phylogeneticand functional analysis of the vertebrate axial musculoskeletalsystem. Traditionally, the evolution of the tail in ray-finnedfishes has been viewed as a transformation from a primitivelyheterocercal (functionally asymmetrical) tail to a homocercaltail in which the axis of rotation during locomotion was vertical,generating a symmetrical thrust. Both phylogenetic and functionalapproaches are used to examine this hypothesis. Major osteologicaland myological features of the tail in ray-finned fishes aremapped onto a phylogeny of ray-finned fishes to discern historicalsequences of morphological change in the axial musculoskeletalsystem. A key event in locomotor evolution was the origin ofthe hypochordal longitudinalis muscle, the only intrinsic caudalmuscle with a line of action at an appreciable angle to thebody axis. This muscle originated prior to the origin of a caudalskeleton bearing both hypaxial and epaxial fin ray supports.The hypochordal muscle is proposed to be a key component ofthe axial musculoskeletal system that allows most fishes tomodulate caudal function and decouples external morphologicalsymmetry from functional symmetry. Experimental data (straingauge recordings from tail bones, and electromyographic recordingsfrom intrinsic and extrinsic caudal muscles) corroborate thisinterpretation and suggest that functional symmetry in the tailof ray-finned fishes is not predictable from skeletal morphologyalone, but depends on the activity of the hypochordal longitudinalismuscle and on locomotor mode. The homocercal teleost tail maythus function asymmetrically.     

Functional Morphology of the Heart in Fishes

The systemic heart of fishes consists of four chambers in series,the sinus venosus, atrium, ventricle, and conus or bulbus. Valvesbetween the chambers and contraction of all chambers exceptthe bulbus maintain a unidirectional blood flow through theheart. The heart is composed of typical vertebrate cardiac muscle,although there may be minor differences in the distributionof spontaneously active cells, the rate and nature of spreadof excitatory waves, and the characteristics of resting andaction potentials between different fish and other vertebrates.Cholinergic fibers innervate the heart, except in hagfish whichhave aneural hearts. Fish hearts lack sympathetic innervation.The level of vagal tone varies considerably, and is affectedby many factors. In some fish the heart is essentially aneural(without vagal tone) during exercise and may resemble an isolatedmammalian ventricle with increased venous return causing increasedcardiac output. There are many mechanisms that could increasevenous return in exercising fish. rß-adrenergic receptorshave been located on the hearts of some fish, and changing levelsof catecholamines may play a role in regulating cardiac activity.Changes in cardiac output in fish are normally associated withlarge changes in stroke volume and small cha-nges in heart rate.     

New Data on Axial Locomotion in Fishes: How Speed Affects Diversity of Kinematics and Motor Patterns

Despite considerable recent progress in understanding the functionof the axial muscles and skeleton in fishes, generalizing fromthese results has been hindered by the great phylogenetic diversityof taxa, the lack of quantitative morphometric data on axialmusculoskeletal structure, and limited analysis of the fullrange of locomotor behaviors exhibited within any one taxon.This paper reviews novel results from our studies of two taxawithin a single monophyletic clade, the sunfish family Centrarchidae.Integrated analyses of lateral displacement, lateral bending,and axial muscle activity reveal widespread effects of swimmingspeed both within a particular mode of swimming and among differentbehavioral modes. The longitudinal position along the body ofthe fish also commonly affects kinematics, muscle activity andthe timing of electromyograms (EMGs) relative to kinematics.EMGs and kinematic events propagate from head to tail for bothsteady and kick and glide swimming. In contrast, during theescape response, the onset of EMGs forms a standing wave pattern,whereas kinematic events are propagated. Several novel featuresof the axial motor pattern are summarized for the kick and glidemode of unsteady swimming. For example, the onset of white fiberEMGs lags significantly behind that of the red fibers at thesame longitudinal position, and red fibers are inactivated athigher unsteady swimming speeds. Muscle activity propagatesposteriorly via the sequential activation of myomeres, but thereare statistically significant differences in the timing of EMGsfrom the contractile tissue opposite a single vertebra. Duringrelatively slow kick and glide swimming, the extreme dorsaland ventral portions of myomeres are not active. Estimates ofthe longitudinal extent of the fish with simultaneous muscleactivity indicate that EMGs from an individual myomere usuallyhave temporal overlap with more than 20 neighboring myomereson the same side of the fish. Consequently, the functional unitsfor axial locomotion of fishes do not correspond simply to theanatomical units of individual myomeres. Rather the in vivomotor pattern is a primary determinant of the functional unitsinvolved in swimming.     

Ecomorphology of Locomotion in Labrid Fishes

The Labridae is an ecologically diverse group of mostly reef associated marine fishes that swim primarily by oscillating their pectoral fins. To generate locomotor thrust, labrids employ the paired pectoral fins in motions that range from a fore-aft rowing stroke to a dorso-ventral flapping stroke. Species that emphasize one or the other behavior are expected to benefit from alternative fin shapes that maximize performance of their primary swimming behavior. We document the diversity of pectoral fin shape in 143 species of labrids from the Great Barrier Reef and the Caribbean. Pectoral fin aspect ratio ranged among species from 1.12 to 4.48 and showed a distribution with two peaks at about 2.0 and 3.0. Higher aspect ratio fins typically had a relatively long leading edge and were narrower distally. Body mass only explained 3% of the variation in fin aspect ratio in spite of four orders of magnitude range and an expectation that the advantages of high aspect ratio fins and flapping motion are greatest at large body sizes. Aspect ratio was correlated with the angle of attachment of the fin on the body (r = 0.65), indicating that the orientation of the pectoral girdle is rotated in high aspect ratio species to enable them to move their fin in a flapping motion. Field measures of routine swimming speed were made in 43 species from the Great Barrier Reef. Multiple regression revealed that fin aspect ratio explained 52% of the variation in size-corrected swimming speed, but the angle of attachment of the pectoral fin only explained an additional 2%. Labrid locomotor diversity appears to be related to a trade-off between efficiency of fast swimming and maneuverability in slow swimming species. Slow swimmers typically swim closer to the reef while fast swimmers dominate the water column and shallow, high-flow habitats. Planktivory was the most common trophic associate with high aspect ratio fins and fast swimming, apparently evolving six times.     

鱼类摄食代谢和运动代谢研究进展

摄食和运动不仅是动物最主要的生理活动,同时也是机体代谢能量消耗的主要过程。相关研究表明鱼类摄食代谢主要由营养物质同化过程的耗能组成,其食物蛋白质同化耗能远低于陆生脊椎动物,而摄入营养物质不平衡可能导致摄食代谢耗能增加;鱼类摄食代谢和运动代谢上可能存在能量消耗与性能维持之间的权衡,且都可能受最大代谢能力限制。鱼类不仅在摄食和运动代谢的相对大小及其他特征上存在差异,而且在摄食和运动代谢竞争上存在不同的模式。从功率分配的角度,研究鱼类摄食和运动代谢特征及其与物种生态习性的关系将成为鱼类能量学研究的重要方向之一。     

Locomotion of Gymnarchus Niloticus： Experiment and Kinematics

In addition to forward undulatory swimming, Gymnarchus niloticus can swim via undulations of the dorsal fin while the body axis remains straight; furthermore, it swims forward and backward in a similar way, which indicates that the undulation of the dorsal fin can simultaneously provide bidirectional propulsive and maneuvering forces with the help of the tail fin. A high-resolution Charge-Coupled Device （CCD） imaging camera system is used to record kinematics of steady swimming as well as maneuvering in G. niloticus. Based on experimental data, this paper discusses the kinematics （cruising speed, wave speed, cycle frequency, amplitude, lateral displacement） of forward as well as backward swimming and maneuvering. During forward swimming, the propulsive force is generated mainly by undulations of the dorsal fin while the body axis remains straight. The kinematic parameters （wave speed, wavelength, cycle frequency, amplitude） have statistically significant correlations with cruising speed. In addition, the yaw at the head is minimal during steady swimming. From experimental data, the maximal lateral displacement of head is not more than 1% of the body length, while the maximal lateral displacement of the whole body is not more than 5% of the body length. Another important feature is that G. niloticus swims backwards using an undulatory mechanism that resembles the forward undulatory swimming mechanism. In backward swimming, the increase of lateral displacement of the head is comparatively significant; the amplitude profiles of the propulsive wave along the dorsal fin are significantly different from those in forward swimming. When G. niloticus does fast maneuvering, its body is first bent into either a C shape or an S shape, then it is rapidly unwound in a travelling wave fashion. It rarely maneuvers without the help of the tail fin and body bending.     

Body Form, Locomotion and Foraging in Aquatic Vertebrates

Four functional categories are denned to embrace the range oflocomotor diversity of aquatic vertebrates; (1) body/caudalfin (BCF) periodic propulsion where locomotor movements repeat,as occurs in cruising and sprint swimming; (2) BCF transientpropulsion where kinematics are brief and non-cylic, as occursin fast-starts and powered turns; (3) median and paired fin(MPF) propulsion, with very diverse fin kinematics, used inslow swimming and precise maneuver; (4) occasional propulsionor "non-swimming." Specialization in any one of these categoriescompromises performance in one or more of the others, therebyreducing locomotor diversity and hence behavioral options. Foodcharacteristics influencing the role of locomotion in searchand capture are; (1) distribution in space and/or time and (2)evasive capabilities. BCF periodic swimmers take food that iswidely dispersed in space/time; BCF transient swimmers consumelocally abundant evasive items and MPF swimmers consume non-evasivefood in structurally complex habitats. Locomotor specialistsunder-utilize smaller food items in exposed habitats. This resourceis exploited by smaller fish, which are locomotor generalistsbecause of predation pressures. For such locomotor generalists,locomotor adaptations for food capture are of diminished importanceand other adaptations such as suction and protrusible jaws infish are common.     

Design and Implementation of Paired Pectoral Fins Locomotion of Labriform Fish Applied to a Fish Robot

In present,there are increasing interests in the research on mechanical and control system of underwater vehicles.Theseongoing research efforts are motivated by more pervasive applications of such vehicles including seabed oil and gas explorations,scientific deep ocean surveys,military purposes,ecological and water environmental studies,and also entertainments.However,the performance of underwater vehicles with screw type propellers is not prospective in terms of its efficiency andmaneuverability.The main weaknesses of this kind of propellers are the production of vortices and sudden generation of thrustforces which make the control of the position and motion difficult.On the other hand,fishes and other aquatic animals are efficient swimmers,posses high maneuverability,are able to followtrajectories,can efficiently stabilize themselves in currents and surges,create less wakes than currently used underwater vehicle,and also have a noiseless propulsion.The fish’s locomotion mechanism is mainly controlled by its caudal fin and paired pectoralfins.They are classified into Body and/or Caudal Fin(BCF)and Median and/or paired Pectoral Fins(MPF).The study of highlyefficient swimming mechanisms of fish can inspire a better underwater vehicles thruster design and its mechanism.There are few studies on underwater vehicles or fish robots using paired pectoral fins as thruster.The work presented in thispaper represents a contribution in this area covering study,design and implementation of locomotion mechanisms of pairedpectoral fins in a fish robot.The performance and viability of the biomimetic method for underwater vehicles are highlightedthrough in-water experiment of a robotic fish.     

Morphology, Velocity, and Intermittent Flight in Birds

Body size, pectoralis composition, aspect ratio of the wing,and forward speed affect the use of intermittent flight in birds.During intermittent non-flapping phases, birds extend theirwings and glide or flex their wings and bound. The pectoralismuscle is active during glides but not during bounds; activityin other primary flight muscles is variable. Mechanical power,altitude, and velocity vary among wingbeats in flapping phases;associated with this variation are changes in neuromuscularrecruitment, wingbeat frequency, amplitude, and gait. Speciesof intermediate body mass (35–158 g) tend to flap-glideat slower speeds and flap-bound at faster speeds, regardlessof the aspect ratio of their wings. Such behavior may reducemechanical power output relative to continuous flapping. Smallerspecies (<20 g) with wings of low aspect ratio may flap-boundat all speeds, yet existing models do not predict an aerodynamicadvantage for the flight style at slow speeds. The behaviorof these species appears to be due to wing shape rather thanpectoralis physiology. As body size increases among species,percent time spent flapping increases, and birds much largerthan 300 g do not flap-bound. This pattern may be explainedby adverse scaling of mass-specific power or lift per unit poweroutput available from flight muscles. The size limit for theability to bound intermittently may be offset somewhat by thescaling of pectoralis composition. The percentage of time spentflapping during intermittent flight also varies according toflight speed.     

Water Striders:The Biomechanics of Water Locomotion and Functional Morphology of the Hydrophobic Surface(Insecta:Hemiptera-Heteroptera)

Water striders are insects living on the water surface, over which they can move very quickly and rarely get wetted. We measured the force of free walking in water striders, using a hair attached to their backs and a 3D strain gauge. The error was calculated by comparing force and data derived from geometry and was estimated as 13%. Females on average were stronger (1.32 mN) than males (0.87 mN), however, the ratio of force to weight was not significantly different. Compared with other lighter species, Aquarius paludum seems stronger, but the ratio of force to weight is actually lower. A. paludum applies about 0.3 mN·cm-1 to 0.4 mN·cm-1 with its mid-legs, thus avoiding penetrating the surface tension layer while propelling itself rapidly over the water surface.We also investigated the external morphology with SEM. The body is covered by effectively two layers of macro-and micro-hairs, which renders them hydrophobic. The setae are long (40 um-60 um) and stiff, being responsible for waterproofing, and the microtrichia are much smaller (<10 um), slender, and flexible, holding a bubble over the body when submerged.     

Morphology and Interrelationships of Primitive Actinopterygian Fishes

SYNOPSIS. The concept of the Actinopterygii as a natural groupof fishes was not generally accepted until early in this century.Ever since, the characterization of the group has been blurredby the problem of cladistian (polypterid) relationships. Froma review of the structure of polypterids and actinopts, it isconcluded that Cladistia are the sistergroup of Recent actinopterygians(Actinopteri), the two groups together comprising the Actinopterygii.Recent chondrosteans are more closely related to higher actinopts(Neopterygii) than to cladistians. The extinct Palaeonisciformesappear to be a paraphyletic group, comprising stem-group actinopterygians(e.g., Cheirolepts), stem-group actinopterans (e.g., Moythomasia)and relatives of higher actinopterans (e.g., Pteroniscus)     

Convergent Evolution of Mechanically Optimal Locomotion in Aquatic Invertebrates and Vertebrates

Examples of animals evolving similar traits despite the absence of that trait in the last common ancestor, such as the wing and camera-type lens eye in vertebrates and invertebrates, are called cases of convergent evolution. Instances of convergent evolution of locomotory patterns that quantitatively agree with the mechanically optimal solution are very rare. Here, we show that, with respect to a very diverse group of aquatic animals, a mechanically optimal method of swimming with elongated fins has evolved independently at least eight times in both vertebrate and invertebrate swimmers across three different phyla. Specifically, if we take the length of an undulation along an animal’s fin during swimming and divide it by the mean amplitude of undulations along the fin length, the result is consistently around twenty. We call this value the optimal specific wavelength (OSW). We show that the OSW maximizes the force generated by the body, which also maximizes swimming speed. We hypothesize a mechanical basis for this optimality and suggest reasons for its repeated emergence through evolution.     

Kinematics,Deformation,and Aerodynamics of a Flexible Flapping Rotary Wing in Hovering Flight

The Flapping Rotary Wing(FRW)is a micro air vehicle wing layout coupling flapping,pitching,and rotating motions.It can gain bencfits in high lift from a fast passive rotating motion,which is tightly related to the passive pitching motion directly caused by wing flexible deformation.Therefore,flexible deformation is crucial for the wing kinematics and aerodynamic performance of an FRW.In this paper,we explored the effct of flexibility on wing kinematics and acrodynamics on the basis of a mechanical FRW model.A photogrammetric method was adopted to capture motion images according to which wing orientations and deformations were reconstructed.Corresponding acrodynamic force was computed using computational fluid dynamic method,and wing kinematics and deformations were used as simulation inputs.The experimental measurements presented the real orientation and deformation pattem of a real FRW.The wing passive deformation of a high-intensity FRW was found to be mainly caused by inertial force,and a linear positive spanwise twist was observed in the FRW.The effects of wing deformation on aerodynamic force production and the underlying mechanism were addressed.Results showed that lift augment,rotating moment enhancement,and power efficiency improvement can be achieved when a wing becomes flexible.Wing spanwise twist mainly accounts for these changes in aerodynamics,and increment in spanwise twist could further contributes to aerodynamic improvement.     

The Morphology and Evolution of the Ear in Actinopterygian Fishes

SYNOPSIS. In this paper we consider various aspects of the anatomyand ultrastructure of the actinopterygian ear and make a numberof suggestions on the possible adaptive significance of thestructural specializations. The focus of the arguments is basedupon the substantial inter-specific variation in teleost auditorysystems as measured anatomically, behaviorally, and physiologically.It is potentially of considerable significance that the majorpoints of inter-specific variation in the teleost ear are associatedwith the gross morphology and ultrastructure of the otolithicorgan most often implicated in sound detection, the sacculus.Analysis of patterns of sacculus ultrastructure has led to theconclusion that there are, in effect, only about five differentsaccular ultrastructural patterns but that these patterns arebroadly found throughout the teleost fishes. Based upon patternsof inter-specific variation in the sacculus and in other aspectsof the ear and more peripheral auditory structures (e.g., swimbladder),it is argued that adaptations encountered in the teleost auditorysystem cannot be used as reliable taxonomic indicators amongfishes. Rather, it is proposed that the teleost auditory systemis quite maleable in the evolutionary sense, and that interspecificsimilarities in many features of the auditory system reflectconvergent evoluuon, rather than phylogenetic affinities. Theactual selective pressures operating in the evoluuon of thefish auditory system are still essentially unknown. In addition,we cannot be certain that similar ear patterns in differentspecies reflect convergent evolution (or common ancestry), orthat conversely, different ear patterns among species reflectdifferences in auditory function.     

The Kinematics and Surface Electromyography Characteristics of Round Kick of Martial Arts Athletes

In order to improve the level of athletes, modern scientific and technological means can be used to understand the characteristics and rules of movement. This study mainly analyzed the whip leg technique of Sanda athletes.Taking ten athletes as an example, the kinematics and surface electromyography(sEMG) data of them were measured, calculated and sorted out when they weredoing the action of round kick. The results showed that the movement completiontime of the first-level athletes was shorter, 0.34 ± 0.33 s. In the stage of turning hipand hitting, the angle of hip joint increased significantly. In the stage of turninghip, there was a significant difference in the angle of hip joint between differentlevels of athletes (p < 0.05), and there was no significant difference in other kinematics characteristics. In the aspect of sEMG, the duration of muscle discharge ofthe first-level athletes was shorter, but there was no significant difference in integrated electromyogram (IEMG) and root mean square (RMS). The experimentalresults reveal the importance of hip joint in the course of round kick and providesome theoretical bases for improving the level of athletes and carrying outtargeted training.     

Aerial Contests, Sexual Selection and Flight Morphology in Solitary Pompilid Wasps

Aerial contest competition has proven to be a challenging phenomenon to interpret in many territorial insects. Because the duels often consist of elaborate and/or high speed ascending maneuvers, the hypothesis that they are settled due to asymmetries in flight performance is intuitively appealing. We evaluated this hypothesis by contrasting differences in known morphological determinants of flight performance between (1) residents vs. non-residents of the territorial wasp, Hemipepsis ustulata and between (2) H. ustulata vs. a non-territorial relative, Pepsis thisbe . In the first contrast, resident male H. ustulata were seen to be larger, and had a tendency for reduced wing loading, but they did not possess greater flight musculature or wing aspect ratios (i.e., more elongated wings) than their non-resident counterparts. In the second contrast, male H. ustulata exhibited clearly greater flight musculature and greater sexual dimorphism in this parameter (males more muscular), and also exhibited a slight tendency for greater wing loading and smaller aspect ratios than males of the patrolling species P. thisbe . Interestingly, although size is linked with territorial success in H. ustulata , males of this species were not larger than male P. thisbe , nor did the former species exhibit greater sexual size dimorphism. These results do not support the hypothesis that the repeated ascending contests of H. ustulata require, and select for, a high acceleration design. However, the observed intraspecific patterns of flight musculature suggest that high acceleration is favored in males of the perching species, perhaps for the ability to intercept passing receptive females.     

Underwater Locomotion in the Desert Locust: Behavioural Choice When Confronted with an Aquatic Barrier

Laboratory studies have shown that adult desert locusts (Schistocerca gregaria) have a swim motor programme, and early field reports describe swimming by bands of hoppers if their route of march is obstructed by a channel of water. However, it is not known whether unconstrained adult locusts will enter water ‘voluntarily’. We found in the laboratory that not only will locusts readily enter water, but that they frequently submerge themselves completely under the water, and cross a water barrier by walking along the bottom. They can stay submerged for up to approximately 9 min, but there is no evidence for significant gas exchange with the water. So far as we are aware the demonstration of this capability for underwater locomotion is a novel finding in this well-studied insect.