Function of the dorsal fin in bluegill sunfish: Motor patterns during four distinct locomotor behaviors

The median fins of fishes are key features of locomotor morphology which function as complex control surfaces during a variety of behaviors. However, very few studies have experimentally assessed median fin function, as most workers focus on axial structures. In particular, the dorsal fin of many teleost fishes possesses both spiny anterior and soft posterior portions which may function separately during locomotion. We analyzed the function of the soft region of the dorsal fin and of the dorsal inclinator (Di) muscles which are the primary muscles responsible for lateral flexion. We used electromyography to measure in vivo Di activity, as well as activity of the red myomeric muscles located at a similar longitudinal position. We quantified motor patterns during four locomotor behaviors: braking and three propulsive behaviors (steady swimming, kick and glide swimming, and C-starts). During the three propulsive swimming behaviors, the timing of Di activity was more similar to that of ipsilateral red myomeric muscle rather than to contralateral myomeric activity, whereas during braking the timing of activity of the Di muscles was similar to that of the contralateral myomeric musculature. During the three propulsive behaviors, when the Di muscles had activity, it was consistent with the function of stiffening the soft dorsal fin to oppose its tendency to bend as a result of the body being swept laterally through the water. In contrast, activity of the Di muscles during braking was consistent with the function of actively flexing the soft dorsal fin towards the side of the fish that had Di activity. Activity of the Di muscles during steady speed swimming was generally sufficient to resist lateral bending of the soft dorsal fin, whereas during high speed kick and glide swimming and C-starts, Di activity was not sufficient to resist the bending caused by resistive forces imposed by the water. Cumulative data from all four behaviors suggest that the Di muscles can be activated independently relative to the myomeric musculature rather than having a single phase relationship with the myomeric muscle common to all of the observed behaviors. © 1996 Wiley-Liss, Inc.     

How swimming fish use slow and fast muscle fibers: implications for models of vertebrate muscle recruitment

We quantified the intensity and duration of electromyograms (emgs) from the red and white axial muscles in five bluegill sunfish (Lepomis macrochirus) which performed three categories of behavior including steady swimming and burst and glide swimming at moderate and rapid speeds. Steady swimming (at 2 lengths/s) involved exclusively red muscle activity (mean posterior emg duration = 95 ms), whereas unsteady swimming utilized red and white fibers with two features of fiber type recruitment previously undescribed for any ectothermic vertebrate locomotor muscle. First, for moderate speed swimming, the timing of red and white activity differed significantly with the average onset time of white lagging behind that of red by approximately 40 ms. The durations of these white emgs were shorter than those of the red emgs (posterior mean = 82 ms) because offset times were effectively synchronous. Second, compared to steady and moderate speed unsteady swimming, the intensity of red activity during rapid unsteady swimming decreased while the intensity of white muscle activity (mean white emg duration = 33 ms) increased. Decreased red activity associated with increased white activity differs from the general pattern of vertebrate muscle recruitment in which faster fiber types are recruited in addition to, but not to the exclusion of, slower fiber types.     

Role of aerobic and anaerobic circular mantle muscle fibers in swimming squid: electromyography

Circular mantle muscle of squids and cuttlefishes consists of distinct zones of aerobic and anaerobic muscle fibers that are thought to have functional roles analogous to red and white muscle in fishes. To test predictions of the functional role of the circular muscle zones during swimming, electromyograms (EMGs) in conjunction with video footage were recorded from brief squid Lolliguncula brevis (5.0-6.8 cm dorsal mantle length, 10.9-18.3 g) swimming in a flume at speeds of 3-27 cm s(-1). In one set of experiments, in which EMGs were recorded from electrodes intersecting both the central anaerobic and peripheral aerobic circular mantle muscles, electrical activity was detected during each mantle contraction at all swimming speeds, and the amplitude and frequency of responses increased with speed. In another set of experiments, in which EMGs were recorded from electrodes placed in the central anaerobic circular muscle fibers alone, electrical activity was not detected during mantle contraction until speeds of about 15 cm s(-1), when EMG activity was sporadic. At speeds greater than 15 cm s(-1), the frequency of central circular muscle activity subsequently increased with swimming speed until maximum speeds of 21-27 cm s(-1), when muscular activity coincided with the majority of mantle contractions. These results indicate that peripheral aerobic circular muscle is used for low, intermediate, and probably high speeds, whereas central anaerobic circular muscle is recruited at intermediate speeds and used progressively more with speed for powerful, unsteady jetting. This is significant because it suggests that there is specialization and efficient use of locomotive muscle in squids.     

Red and white muscle activity and kinematics of the escape response of the bluegill sunfish during swimming

Summary We quantified midline kinematics with synchronized electromyograms (emgs) from the red and white muscles on both sides of bluegill sunfish (Lepomis macrochirus) during escape behaviors which were elicited from fish both at a standstill and during steady speed swimming. Analyses of variance determined whether or not kinematic and emg variables differed significantly between muscle fiber types, among longitudinal positions, and between swimming versus standstill trials.At a given longitudinal location, both the red and white muscle were usually activated synchronously during both stages of the escape behavior. Stage 1 emg onsets were synchronous; however, the mean durations of stage 1 emgs showed a significant increase posteriorly from about 11 to 15 ms. Stage 2 emgs had significant posterior propagation, but the duration of the stage 2 emgs was constant (17 ms). Posterior emgs from both stages occurred during lengthening of the contractile tissue (as indicated by lateral bending). Steady swimming activity was confined to red muscle bursts which were propagated posteriorly and had significant posterior decrease in duration from about 50% to 37% of a cycle. Fish performed escape responses during all phases of the steady swimming motor pattern. All kinematic events were propagated posteriorly. Furthermore, no distinct kinematic event corresponded to the time intervals of the stage 1 and 2 emgs. The rate of propagation of kinematic events was always slower than that of the muscle activity. The phase relationship between lateral displacement and lateral bending also changed along the length of the fish. Escape responses performed during swimming averaged smaller amplitudes of stage 2 posterior lateral displacement; however, most other kinematic and emg variables did not vary significantly between these two treatments.Abbreviations A angle of lateral flexion (bending) of midline at a single point in time - A1, A2 change in A from T₀ to T₁ and from T₁ to T₂ - AMX maximal lateral flexion concave towards the side of the stage 1 emg - AMXR equals AMX minus A at T₀ - AT1, AT2 lateral flexion at T₁ and T₂ - DUR1, DUR2 durations of stage 1 and stage 2 emgs - emg electromyogram - ON2 onset time of stage 2 emg - RELDUR relative duration of steady swimming emg - T₀, T₁, T₂ times of stage 1 emg onset, latest stage 1 emg offset and latest stage 2 emg offset standardized such that T₀ = 0 - TAMX, TAMN, TYMX times of maximal lateral flexion, no lateral flexion and maximum lateral displacement - Y1, Y2 amounts of lateral displacement from T₀ to T₁ and from T₁ to T₂ - YMXR relative amount of lateral displacement from T₀ to TYMX     

Evolution of behavior and neural control of the fast-start escape response

The fast-start startle behavior is the primary mechanism of rapid escape in fishes and is a model system for examining neural circuit design and musculoskeletal function. To develop a dataset for evolutionary analysis of the startle response, the kinematics and muscle activity patterns of the fast-start were analyzed for four fish species at key branches in the phylogeny of vertebrates. Three of these species (Polypterus palmas, Lepisosteus osseus, and Amia calva) represent the base of the actinopterygian radiation. A fourth species (Oncorhynchus mykiss) provided data for a species in the central region of the teleost phylogeny. Using these data, we explored the evolution of this behavior within the phylogeny of vertebrates. To test the hypothesis that startle features are evolutionarily conservative, the variability of motor patterns and kinematics in fast-starts was described. Results show that the evolution of the startle behavior in fishes, and more broadly among vertebrates, is not conservative. The fast-start has undergone substantial change in suites of kinematics and electromyogram features, including the presence of either a one- or a two-stage kinematic response and change in the extent of bilateral muscle activity. Comparative methods were used to test the evolutionary hypothesis that changes in motor control are correlated with key differences in the kinematics and behavior of the fast-start. Significant evolutionary correlations were found between several motor pattern and behavioral characters. These results suggest that the startle neural circuit itself is not conservative. By tracing the evolution of motor pattern and kinematics on a phylogeny, it is shown that major changes in the neural circuit of the startle behavior occur at several levels in the phylogeny of vertebrates.     

Morphology and mechanics of myosepta in a swimming salamander (Siren lacertina)

In contrast to the complex, three-dimensional shape of myomeres in teleost fishes, the lateral hypaxial muscles of salamanders are nearly planar and their myosepta run in a roughly straight line from mid-lateral to mid-ventral. We used this relatively simple system as the basis for a mathematical model of segmented musculature. Model results highlight the importance of the mechanics of myosepta in determining the shortening characteristics of a muscle segment. We used sonomicrometry to measure the longitudinal deformation of myomeres and the dorsoventral deformation of myosepta in a swimming salamander (Siren lacertina). Sonomicrometry results show that the myosepta allow some dorsoventral lengthening, indicating an amplification of myomere shortening that is greater than that produced by muscle fiber angle alone (10% muscle fiber shortening produces 28.7% myomere shortening). Polarized light and DIC microscopy of isolated hypaxial myosepta revealed that the collagen fiber orientation in hypaxial myomeres is primarily mediolateral. The mediolateral collagen fiber orientation, combined with our finding that the hypaxial myosepta lengthen dorsoventrally during swimming, suggests that one possible function of hypaxial myosepta in S. lacertina is to increase the strain amplification of the muscle fibers by reducing the mediolateral bulging of the myomeres and redirecting the bulging toward the dorsoventral direction.     

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.     

Lungfish Axial Muscle Function and the Vertebrate Water to Land Transition

The role of axial form and function during the vertebrate water to land transition is poorly understood, in part because patterns of axial movement lack morphological correlates. The few studies available from elongate, semi-aquatic vertebrates suggest that moving on land may be powered simply from modifications of generalized swimming axial motor patterns and kinematics. Lungfish are an ideal group to study the role of axial function in terrestrial locomotion as they are the sister taxon to tetrapods and regularly move on land. Here we use electromyography and high-speed video to test whether lungfish moving on land use axial muscles similar to undulatory swimming or demonstrate novelty. We compared terrestrial lungfish data to data from lungfish swimming in different viscosities as well as to salamander locomotion. The terrestrial locomotion of lungfish involved substantial activity in the trunk muscles but almost no tail activity. Unlike other elongate vertebrates, lungfish moved on land with a standing wave pattern of axial muscle activity that closely resembled the pattern observed in terrestrially locomoting salamanders. The similarity in axial motor pattern in salamanders and lungfish suggests that some aspects of neuromuscular control for the axial movements involved in terrestrial locomotion were present before derived appendicular structures.     

Shape variation in a benthic stream fish across flow regimes

Evolution of fish body shapes in flowing and non-flowing waters have been examined for several species. Flowing water can select for fish body shapes that increase steady swimming efficiency, whereas non-flowing water can favor shapes that increase unsteady swimming efficiency. Benthic stream fishes often use areas near the substrate that exhibit reduced or turbulent flow, thus it is unclear which swimming forms would be favored in such environments, and how shape might change across flow regimes. To test the relationship between fish body shape and flow regime in a benthic stream fish, we used geometric morphometric techniques to characterize lateral body shape in mountain sucker (Catostomus platyrhynchus) across flow rates, using stream gradient as an indicator of stream flow. Mountain suckers from low-flow environments were more streamlined, consistent with steady swimming body shapes, whereas mountain suckers from high flows had deeper bodies, consistent with unsteady swimming body shapes. In addition, smaller individuals tended to have more robust body shapes. These patterns are opposite to those predicted for stream fishes in the mid-water column. The benthic stream environment represents a distinct selective environment for fish shape that does not appear to conform to the simple dichotomy of flowing versus non-flowing water.     

Muscle Dynamics in Fish During Steady Swimming

SYNOPSIS. Recent research in fish locomotion has been dominatedby an interest in the dynamic mechanical properties of the swimmingmusculature. Prior observations have indicated that waves ofmuscle activation travel along the body of an undulating fishfaster than the resulting waves of muscular contraction, suggestingthat the phase relation between the muscle strain cycle andits activation must vary along the body. Since this phase relationis critical in determining how the muscle performs in cycliccontractions, the possibility has emerged that dynamic musclefunction may change with axial position in swimming fish. Quantificationof muscle contractile properties in cyclic contractions relieson in vitro experiments using strain and activation data collectedin vivo. In this paper we discuss the relation between theseparameters and body kinematics. Using videoradiographic datafrom swimming mackerel we demonstrate that red muscle straincan be accurately predicted from midline curvature but not fromlateral displacement. Electromyographic recordings show neuronalactivation patterns that are consistent with red muscle performingnet positive work at all axial positions. The relatively constantcross-section of red muscle along much of the body suggeststhat positive power for swimming is generated fairly uniformlyalong the length of the fish.     

Comparative morphology of the myomeres and axial skeleton in four genera of centrarchid fishes

We used X-rays and dissection of myotomes to quantify the axial morphology of four species of centrarchid fishes (Micropterus salmoides, Ambloplites rupestris, Pomoxis nigromaculatus, and Lepomis macrochirus). Proceeding from dorsal to ventral, we designated the two epaxial and two hypaxial protions of the myomeres AB, BC, CD, and DE, respectively. For each of 11 myomeres, spaced at 10% increments along the length of the fish, a total of 14 variables described the length and orientation of each portion, the dorsalventral symmetry, and the overall height and longitudinal span of the entire myomere. Nine variables described the lengths, orientation, and symmetry of the vertebral centra, neural and hemal spines, and ribs. Analysis of variance revealed that, with one exception, all 23 morphological variables varied significantly both among species and among longitudinal locations within a species. However, the extent of longitudinal and interspecific variance differed considerably among different variables. Maximal myomeric height ranged from about 45% of the standard length (SL) in Lepomis to 27% SL in Micropterus. Longitudinal and interspecific increases in overall height of the trunk myomeres resulted primarily from greater lengths of CD. Compared to other portions of the myomere, the length of BC was most conservative both longitudinally and interspecifically. Dorsal-ventral symmetry of the myomeres and axial skeleton was greater in the caudal region than in the trunk in all species, and the myomeric morphology diverged least among species in the posterior caudal region. The overall longitudinal span of superficial myomeric landmarks varied from 6% to 18% SL, and, including the deep portions of the myomeres, the longitudinal span varied from about 7 to 10 vertebrae. Within each of the species, myomeric and skeletal variables were often not significantly correlated, but for the pooled data of all species there were usually highly significant correlations between myomeric and skeletal morphology. For example, strong correlations existed between BC and the underlying neural spines, and between CD and the underlying ribs and hemal spines. In contrast, the longitudinal spans of entire myomeres and underlying axial skeletal segments were only weakly associated. © 1994 Wiley-Liss, Inc.     

Muscle activity in steady swimming scup, Stenotomus chrysops, varies with fiber type and body position

The red and pink aerobic muscle fibers are used to power steady swimming in fishes. We examined red and pink muscle recruitment and function during swimming in scup, Stenotomus chrysops, through electromyography and high-speed ciné. Computer analysis of electromyograms (EMGs) allowed determination of initial speed of muscle recruitment and duty cycle and phase of muscle electromyographic activity for both fiber types. This analysis was carried out for three longitudinal positions over a range of swimming speeds. Fiber type and longitudinal position both affected swimming speed of initial recruitment. Posterior muscle is recruited at the lowest swimming speed, whereas more anterior muscle is not initially recruited until higher speeds. At more anterior positions, the initial recruitment of pink muscle occurs at a higher swimming speed than the recruitment of red muscle. The duty cycle of pink muscle EMG activity is significantly shorter than that of red muscle, reflecting a difference in the onset time of activation during each cycle of length change: pink muscle onset time follows that of red. The different patterns of usage of red and pink muscle reflect differences in their contraction kinetics. Because pink muscle generates force more rapidly than red muscle, it can be activated later in each tailbeat cycle. Pink muscle is used to augment red muscle power production at higher swimming speeds, allowing a higher aerobically based steady swimming speed than that possible by red muscle alone.     

Evolutionary transformations of myoseptal tendons in gnathostomes

Axial undulations in fishes are powered by a series of three-dimensionally folded myomeres separated by sheets of connective tissue, the myosepta. Myosepta have been hypothesized to function as transmitters of muscular forces to axial structures during swimming, but the difficulty of studying these delicate complex structures has precluded a more complete understanding of myoseptal mechanics. We have developed a new combination of techniques for visualizing the three-dimensional morphology of myosepta, and here we present their collagen-fibre architecture based on examination of 62 species representing all of the major clades of notochordates. In all gnathostome fishes, each myoseptum bears a set of six specifically arranged tendons. Because these tendons are not present outside the gnathostomes (i.e. they are absent from lampreys, hagfishes and lancelets), they represent evolutionary novelties of the gnathostome ancestor. This arrangement has remained unchanged throughout 400 Myr of gnathostome evolution, changing only on the transition to land. The high uniformity of myoseptal architecture in gnathostome fishes indicates functional significance and may be a key to understanding general principles of fish swimming mechanics. In the design of future experiments or biomechanical models, myosepta have to be regarded as tendons that can distribute forces in specific directions.     

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.     

Radio-transmitted electromyogram signals as indicators of physical activity in Atlantic salmon

Surgical methods developed to implant EMG (electromyogram) transmitters in Atlantic salmon Salmo salar were tested to calibrate electromyograms from axial red musculature to swimming speed in a swim speed chamber, and to compare electromyograms of fish from two stocks (Lone and Imsa). Ten Lone and eight Imsa salmon were equipped with internal EMG transmitters. Surgical procedures were acceptable, with 100% survival of all implanted fish during the study. It was possible to calibrate EMG pulse intervals to swimming speed in 14 of the 18 salmon run in the swim speed chamber ( r²= 0·35-0·76 for individuals, 0·63 for pooled data). Individuals differed in their EMG resting levels (EMGs recorded at 0·5 ms⁻¹), and so higher correlations were obtained between swimming speed and an activity index (EMG pulse intervals at different speeds/EMG resting levels) (pooled data, r² =0·75). The linear relationship between swimming speed and EMG pulse intervals differed significantly between the two stocks ( P <0·05). This successful calibration of EMGs to swimming speed opens the possibility of calibrating EMGs to oxygen consumption and the measurement of the metabolic costs of activity in field experiments.     

Functional morphology of ventral tail bending and prehensile abilities of the seahorse,Hippocampus kuda

Unlike most teleosts, the seahorse (genus Hippocampus) is able to bend its tail ventrally, uses its tail in a postural role as a grasping and holding appendage, and possesses heavy body plates instead of scales. To investigate seahorse axial bending mechanisms and the role of plating in those mechanisms, observations were made on seahorses curling their tails ventrally and holding a support and components of the mechanical system used for axial bending, including dermal plates, vertebrae, and axial muscles, were examined. Anatomical modifications involved in ventral tail bending include hypertrophy of the ventral region of the hypaxial muscle, ventrolateral attachment of the myomeres to plates, and modification of the infracarinalis posterior muscles so that they act in axial bending rather than in fin movement as has previously been hypothesized (Harder, '75) for other fishes. Modifications for prehension include the presence of fibers histochemically characterized as tonic in the median ventral muscles (the modified infracarinalis muscle) and in portions of the myomeres. Dermal plates are an important part of the force transmission system used in seahorse tail bending. They transmit forces from the hypaxial myomeres to bend the tail both laterally and ventrally. This study expands our understanding of axial bending in fishes by examining extreme modifications of the musculoskeletal system associated with the evolution of unique functional capabilities within teleosts. © 1996 Wiley-Liss, Inc.     

Myoseptal architecture of sarcopterygian fishes and salamanders with special reference to Ambystoma mexicanum

During axial undulatory swimming in fishes and salamanders muscular forces are transmitted to the vertebral axis and to the tail. One of the major components of force transmission is the myoseptal system. The structure of this system is well known in actinopterygian fishes, but has never been addressed in sarcopterygian fishes or salamanders. In this study we describe the spatial arrangement and collagen fiber architecture of myosepta in Latimeria, two dipnoans, and three salamanders in order to gain insight into function and evolution of the myoseptal system in these groups. Salamander myosepta lack prominent cones, and consist of homogenously distributed collagen fibers of various orientations that never form distinct tendons. Fiber orientations are difficult to homologize with those of fish myosepta. The myosepta of Latimeria and dipnoans (Protopterus and Neoceratodus) illustrate that major changes in architecture occurred in the sarcopterygian clade (loss of horizontal septum), in the rhipidistian (dipnoans + tetrapods) clade (loss of epineural and epipleural tendon), and in tetrapods (loss of lateral tendons and myoseptal folding). When compared to fishes, the myosepta of wholly aquatic salamanders (Ambystoma mexicanum, Amphiuma tridactylum, Necturus maculosus) do not have the lateral tendons we suppose serve to transfer muscular forces posteriorly. We propose that alternative structures (most conspicuously present in Ambystoma) perform this function: posteriorly the relative amount of connective tissue increases considerably, and myosepta are disintegrated to horizontal lamellae of connective tissue. The structures thought to be involved in modulation of body stiffness in fishes during swimming are also absent in salamanders. Our data also have implications for the hypothesis that salamander hypaxial myosepta are designed to increase shortening amplification of the hypaxial muscle fibers. The posterior hypaxial myosepta of all three salamander species possess only mediolaterally directed collagen fibers, which would indeed amplify the shortening of the associated muscle.     

Oxygen consumption of active rainbow trout, Salmo gairdneri Richardson, derived from electromyograms obtained by radiotelemetry

A radiotelemetry apparatus is described for sensing and transmitting electromyograms (EMGs) from free-swimming fish. EMGs are recorded from the epaxial muscles of adult rainbow trout during periods in spontaneous (= routine) activity, and forced-swim, respirometers. When such EMG records are integrated, subjected to spectral analysis, and computer-averaged, the EMG values (in μV) are highly correlated with the fish oxygen consumption during the activity periods. However, there is a marked difference between the regression slopes for oxygen v . EMG value for the data from the spontaneous, and forced-swim, respirometers; the former slope is the steeper. The probable explanation of this phenomenon is that whereas in forced swims the epaxial myomeres are responsible for most of the activity of the fish, in spontaneous activity other muscle systems (e.g. of the lateral, dorsal and ventral fins) come to account for a greater relative proportion of body movement. The difference in slope, although great, is evidently a regular phenomenon. The shift from one regression to the other occurs at a fairly precise epaxial EMG value ( c . 5 μV). This suggests that the laboratory calibration of EMG value in terms of oxygen consumption can be utilized in the wild so that EMG records from free-swimming fish, fitted with telemetry packages can be used to deduce oxygen consumption attributable to activity. It also appears that such records can be used as a guide to the type of activity of the fish, i.e. desultory movements or free cruising.     

Kinematics and hydrodynamics of linear acceleration in eels, Anguilla rostrata

The kinematics and hydrodynamics of routine linear accelerations were studied in American eels, Anguilla rostrata, using high-speed video and particle image velocimetry. Eels were examined both during steady swimming at speeds from 0.6 to 1.9 body lengths (L) per second and during accelerations from -1.4 to 1.3 L s(-2). Multiple regression of the acceleration and steady swimming speed on the body kinematics suggests that eels primarily change their tail-tip velocity during acceleration. By contrast, the best predictor of steady swimming speed is body wave speed, keeping tail-tip velocity an approximately constant fraction of the swimming velocity. Thus, during steady swimming, Strouhal number does not vary with speed, remaining close to 0.32, but during acceleration, it deviates from the steady value. The kinematic changes during acceleration are indicated hydrodynamically by axial fluid momentum in the wake. During steady swimming, the wake consists of lateral jets of fluid and has minimal net axial momentum, which reflects a balance between thrust and drag. During acceleration, those jets rotate to point downstream, adding axial momentum to the fluid. The amount of added momentum correlates with the acceleration, but is greater than the necessary inertial force by 2.8+/-0.6 times, indicating a substantial acceleration reaction.