Functional morphology of the muscles in Philodina sp. (Rotifera: Bdelloidea)

Whole-mounts of Philodina sp., a bdelloid rotifer, were stained with fluorescent-labeled phalloidin to visualize the musculature. Several different muscle types were identified including incomplete circular bands, coronal retractors and foot retractors. Based on the position of the larger muscle bands in the body wall, their function during creeping locomotion and tun formation was inferred. Bdelloid creeping begins with the contraction of incomplete circular muscle bands against the hydrostatic pseudocoel, resulting in an anterior elongation of the body. One or more sets of ventral longitudinal muscles then contract bringing the rostrum into contact with the substrate, where it presumably attaches via adhesive glands. Different sets of ventral longitudinal muscles, foot and trunk retractors, function to pull the body forward. These same longitudinal muscle sets are also used in `tun' formation, in which the head and foot are withdrawn into the body. Three sets of longitudinal muscles supply the head region (anterior head segments) and function in withdrawal of the corona and rostrum. Two additional pairs of longitudinal muscles function to retract the anterior trunk segments immediately behind the head, and approximately five sets of longitudinal retractors are involved in the withdrawal of the foot and posterior toes. To achieve a greater understanding of rotifer behavior, it is important to elucidate the structural complexity of body wall muscles in rotifers. The utility of fluorescently-labeled phalloidin for the visualization of these muscles is discussed and placed in the context of rotifer functional morphology.     

Locomotion of free-swimming ghost knifefish: anal fin kinematics during four behaviors

The maneuverability demonstrated by the weakly electric ghost knifefish (Apteronotus albifrons) is a result of its highly flexible ribbon-like anal fin, which extends nearly three-quarters the length of its body and is composed of approximately 150 individual fin rays. To understand how movement of the anal fin controls locomotion we examined kinematics of the whole fin, as well as selected individual fin rays, during four locomotor behaviors executed by free-swimming ghost knifefish: forward swimming, backward swimming, heave (vertical) motion, and hovering. We used high-speed video (1000 fps) to examine the motion of the entire anal fin and we measured the three-dimensional curvature of four adjacent fin rays in the middle of the fin during each behavior to determine how individual fin rays bend along their length during swimming. Canonical discriminant analysis separated all four behaviors on anal fin kinematic variables and showed that forward and backward swimming behaviors contrasted the most: forward behaviors exhibited a large anterior wavelength and posterior amplitude while during backward locomotion the anal fin exhibited both a large posterior wavelength and anterior amplitude. Heave and hover behaviors were defined by similar kinematic variables; however, for each variable, the mean values for heave motions were generally greater than for hovering. Individual fin rays in the middle of the anal fin curved substantially along their length during swimming, and the magnitude of this curvature was nearly twice the previously measured maximum curvature for ray-finned fish fin rays during locomotion. Fin rays were often curved into the direction of motion, indicating active control of fin ray curvature, and not just passive bending in response to fluid loading.     

Muscular mechanisms of snake locomotion: an electromyographic study of lateral undulation of the Florida banded water snake (Nerodia fasciata) and the yellow rat snake (Elaphe obsoleta)

Electromyography and cinematography were used to determine the activity of epaxial muscles of colubrid snakes during terrestrial and aquatic lateral undulatory locomotion. In both types of lateral undulation, at a given longitudinal position, segments of three muscles (Mm. semispinalis-spinalis, longissimus dorsi, and iliocostalis) usually show synchronous activity. Muscle activity propagates posteriorly and generally is unilateral. With each muscle, large numbers of adjacent segments (30 to 100) show simultaneous activity. Terrestrial and aquatic undulation differ in two major respects. (1) During terrestrial undulation, muscle activity in a particular region begins when that portion of the body has reached maximal convex flexion and ends when it is maximally concave; this phase relation is uniform along the entire snake. During swimming, however, muscle activity passes posteriorly faster than the wave of vertebral flexion, causing the relation of muscle activity to flexion to change along the length of the snake. (2) In the terrestrial mode, the block of active muscle segments remains approximately constant in size as it passes down the snake, whereas during swimming the number of adjacent active muscle segments increases posteriorly. Despite the fact that Elaphe obsoleta has nearly twice as many body vertebrate as Nerodia fasciata (240 vs. 125), the only difference observed in the swimming of these two species is that a larger number of adjacent muscle segments is simultaneously active in comparable regions of Elaphe obsoleta than in Nerodia fasciata.     

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.     

Power isn't everything: muscle function and energetic costs during steady swimming in Atlantic cod (Gadus morhua)

Power produced by red myotomal muscles of fish during cruise swimming appears seldom maximized, so we sought to investigate whether economy may impact or dominate muscle function. We measured cost of transport (COT) using oxygen consumption and the strain trajectories and electromyographic activity of red muscle measured at anterior (ANT) and posterior (POST) locations while Atlantic cod (Gadus morhua) swam steadily at speeds between 0.3 and 1.0 body lengths (BL) s(-1). We then measured the power produced by isolated segments of red muscle when activated either as in the swimming cod or such that maximal net power was produced. Patterns of activation during swimming were not optimal for power output and were highly variable between tail beats, particularly at the ANT location and at slow swim speeds. Muscle strain amplitude did not increase until swimming speed reached 0.9 (ANT) versus 0.5 (POST) BL s(-1). These limited power to only 53% (ANT) and 71% (POST) of maximum at slower swim speeds and to 70%-80% of maximum at high swim speeds. COT (resting metabolism subtracted) was minimal at the slowest swim speed, surprisingly, where power was most impaired by activation and strain. Thus, production of powered forces for maneuverability/stability appeared to greatly impact red muscle function during cruise swimming in cod, particularly at slow speeds and in ANT muscle.     

The burrowing of Priapulus caudatus

An account is given of the way in which Priapulus caudatus burrows in the muddy sea-bed in which it lives. Three phases are distinguishable in the muscular activity which is responsible for locomotion. During the first phase the animal is able to feed and defaecate, during the second the proboscis becomes invaginated, and during the third the animal moves forward. The power for locomotion is provided by contraction of the longitudinal and circular muscles of the body wall, not, as has been suggested previously, by the retractor muscles of the praesoma. Invagination of the proboscis is apparently stimulated by the arrival of a wave of contraction in the body wall musculature, propagated from the trunk.
In general the animal burrows in a way common for soft-bodied animals; the anterior and posterior extremities acting in turn as "terminal" and "penetration" anchors in the substratum. The muscular activities of the larva are limited by the presence of a lorica which encases the trunk, and the animal's powers of movement at this stage are very restricted.     

Flexibility in the patterning and control of axial locomotor networks in lamprey

In lower vertebrates, locomotor burst generators for axial muscles generally produce unitary bursts that alternate between the two sides of the body. In lamprey, a lower vertebrate, locomotor activity in the axial ventral roots of the isolated spinal cord can exhibit flexibility in the timings of bursts to dorsally-located myotomal muscle fibers versus ventrally-located myotomal muscle fibers. These episodes of decreased synchrony can occur spontaneously, especially in the rostral spinal cord where the propagating body waves of swimming originate. Application of serotonin, an endogenous spinal neurotransmitter known to presynaptically inhibit excitatory synapses in lamprey, can promote decreased synchrony of dorsal-ventral bursting. These observations suggest the possible existence of dorsal and ventral locomotor networks with modifiable coupling strength between them. Intracellular recordings of motoneurons during locomotor activity provide some support for this model. Pairs of motoneurons innervating myotomal muscle fibers of similar ipsilateral dorsoventral location tend to have higher correlations of fast synaptic activity during fictive locomotion than do pairs of motoneurons innervating myotomes of different ipsilateral dorsoventral locations, suggesting their control by different populations of premotor interneurons. Further, these different motoneuron pools receive different patterns of excitatory and inhibitory inputs from individual reticulospinal neurons, conveyed in part by different sets of premotor interneurons. Perhaps, then, the locomotor network of the lamprey is not simply a unitary burst generator on each side of the spinal cord that activates all ipsilateral body muscles simultaneously. Instead, the burst generator on each side may comprise at least two coupled burst generators, one controlling motoneurons innervating dorsal body muscles and one controlling motoneurons innervating ventral body muscles. The coupling strength between these two ipsilateral burst generators may be modifiable and weakening when greater swimming maneuverability is required. Variable coupling of intrasegmental burst generators in the lamprey may be a precursor to the variable coupling of burst generators observed in the control of locomotion in the joints of limbed vertebrates.     

The swimming endurance of threespine sticklebacks, Gasterosteus aculeatus L., from the Afon Rheidol, Wales

The endurance of threespine sticklebacks, Gasterosteus aculeatus , swimming with pectoral fin locomotion at 20° C in a laboratory flume was measured. Each trial lasted a maximum of 480 min. At a speed of 4 body lengths per sec (L s⁻¹) all fish were still swimming at the end of the trial, but endurance decreased at higher speeds. At speeds of 5 or 6 L s⁻¹ (20–30 cm s⁻¹) a few fish still maintained labriform locomotion for the 480 min. However, at a speed of 7 L s⁻¹ all fish furled their pectoral fins and used body and caudal fin propulsion but fatigued rapidly. During sustained swimming, fish could cover distances of 6 km or more. No significant differences between males and females were found.     

Behavioral plasticity and central regeneration of locomotor reflexes in the freshwater oligochaete, Lumbriculus variegatus. II: Ablation studies

Abstract. After 8–10 segments of posterior ventral nerve cord were ablated in Lumbriculus variegatus , touch-evoked locomotor responses were evident both in segments anterior and posterior to the ablation site. However, responses in these two regions were independent and uncoupled. During recovery, four outcomes were observed at the ablation site: (Group 1) recovery of normal functions with no growth of new segments; (Group 2) formation of a laterally protruding, multi-segmented, ectopic head; (Group 3) formation of a laterally protruding, amorphous, and multi-segmented outgrowth; and (Group 4) segmental autotomy. In Groups 1 and 2, touch-evoked swimming and body reversal were studied. In addition, sensory fields and conduction properties of giant nerve fibers were examined near the ablation site. In some Group 1 worms, clear-cut behavioral and electrical signs of recovery and reconnection were seen by 3 d after ablation. By 8 d, all worms had recovered and exhibited response patterns comparable to those of normal worms. In Group 2 worms, with an ectopic head, segments posterior to the ablation (together with those in the ectopic head), exhibited touch-evoked swimming and body reversal responses resembling those of a complete worm. Segments anterior to the ectopic head were independently capable of locomotor responses. Medial and lateral giant fiber sensory fields in worms with ectopic heads reflected a pattern expected for two worms. Thus, through apparent morphallactic reorganization, a medial giant fiber sensory field emerged which included the ectopic head and 10–15 adjacent posterior segments. In contrast, electrical recordings showed longitudinal through-conduction of giant fiber spikes, across the ablation site. Histological examination revealed that the giant nerve fibers in the ectopic head were complexly interconnected with those in the main body axis.     

The relationship between body size and the field potentials generated by swimming crayfish

Aquatic animals generate electrical field potentials which may be monitored by predators or conspecifics. Many crustaceans use rapid, forceful contractions of the flexor and extensor muscles to curl and extend their abdomens during swimming in escape and locomotion. When crayfish swim they generate electrical field potentials that can be recorded by electrodes nearby in the water. In general, it is reasonable to assume that larger bodied crayfish will generate signals of greater amplitude because they have larger muscles. It is not known, however, how activity in particular muscles and nerves combines to produce the compound electrical waveform recorded during swimming. We therefore investigated the relationship between abdominal muscle, body size and the amplitude of nearby tailflip potentials in the freshwater crayfish (Cherax destructor). We found that amplitude was correlated positively with abdominal muscle mass. The mean amplitude recorded from the five smallest and five largest individuals differed by 440 microV, a difference sufficiently large to be of significance to predators and co-inhabitants in the wild.     

Resolving Shifting Patterns of Muscle Energy Use in Swimming Fish

Muscle metabolism dominates the energy costs of locomotion. Although in vivo measures of muscle strain, activity and force can indicate mechanical function, similar muscle-level measures of energy use are challenging to obtain. Without this information locomotor systems are essentially a black box in terms of the distribution of metabolic energy. Although in situ measurements of muscle metabolism are not practical in multiple muscles, the rate of blood flow to skeletal muscle tissue can be used as a proxy for aerobic metabolism, allowing the cost of particular muscle functions to be estimated. Axial, undulatory swimming is one of the most common modes of vertebrate locomotion. In fish, segmented myotomal muscles are the primary power source, driving undulations of the body axis that transfer momentum to the water. Multiple fins and the associated fin muscles also contribute to thrust production, and stabilization and control of the swimming trajectory. We have used blood flow tracers in swimming rainbow trout (Oncorhynchus mykiss) to estimate the regional distribution of energy use across the myotomal and fin muscle groups to reveal the functional distribution of metabolic energy use within a swimming animal for the first time. Energy use by the myotomal muscle increased with speed to meet thrust requirements, particularly in posterior myotomes where muscle power outputs are greatest. At low speeds, there was high fin muscle energy use, consistent with active stability control. As speed increased, and fins were adducted, overall fin muscle energy use declined, except in the caudal fin muscles where active fin stiffening is required to maintain power transfer to the wake. The present data were obtained under steady-state conditions which rarely apply in natural, physical environments. This approach also has potential to reveal the mechanical factors that underlie changes in locomotor cost associated with movement through unsteady flow regimes.     

Neural Control of Swimming in the Leech

Leeches swim by undulating; they alternately form crests thentroughs at their anterior end and move Them backward, therebyproducing forward thrust. These movements are accomplished byalternating contractions of dorsal and ventral longitudinalmuscles in each of the 21 body segments. These contractionsare caused by bursts of impulses in groups of excitatory andinhibitory motor neurons. Connections among motor neurons helpto coordinate these bursts: synergistic muscle excitors areelectrotonically coupled, which aids in keeping their burstsnearly synchronous; muscle inhibitors also inhibit the excitorsto the same muscles, and it is this inhibition which keeps theexcitors from being tonically active during swimming. Neuronssensitive to either dorsal or ventral body wall stretch producereciprocal stretch reflexes to the muscle excitors, probablyvia the inhibitors. That these stretch reflexes may be involvedin generating the periodic bursts is supported by the resultsof both behavioral and electrophysiological experiments.     

Integrative analysis of polychaete ontogeny: cell proliferation patterns and myogenesis in trochophore larvae of Sabellaria alveolata

SUMMARY Aspects of muscle development are still widely neglected in studies on invertebrate ontogeny, which is probably at least partly due to the inherent complexity of animal myoanatomical bodyplans. This has resulted in significant gaps in our understanding of the evolutionary and ontogenetic origin of this crucial mesoderm-derived organ system, particularly in indirect developing representatives of the Lophotrochozoa. Here, we document the temporal and spatial patterns of muscle formation and cell proliferation in the polychaete Sabellaria alveolata during planktotrophic larval development in order to assess the presumed generation of muscle units and segments from a posterior growth zone. In addition, we address the question whether the three primary segments differ in their mode of myogenesis from the subsequently forming segments. We found that in the first three segments the ventral transverse muscles differentiate from anterior to posterior, whereas the ventral oblique muscles develop simultaneously. Hence, subsequent and simultaneous developmental processes of specific muscle groups are possibly regulated in different ways, thus emphasizing the plasticity of the formation of metamerically arranged organ systems in polychaetes. The occurrence of three clusters of proliferating cells in the trunk region of the metatrochophore indicates synchronous subdivision of the larval mesoderm in the first three segments. Assuming a polychaete-like ancestor at the base of the annelid tree, comparative analysis suggests that the bodywall of the last common annelid ancestor might have been devoid of circular muscles and consisted of four separate longitudinal muscle strands that develop from anterior to posterior.     

Functional morphology of muscles in Tetranchyroderma papii (Gastrotricha)

Movement in gastrotrichs is powered by an interaction of ventral cilia and muscles. In interstitial gastrotrichs, movement among sand grains often requires the additional use of adhesive tubules that allow for behaviors such as escape responses and changes in body position. In this study, we describe orientations and possible mechanical actions of muscles during locomotion in the gastrotrich, Tetranchyroderma papii (Macrodasyida). Fluorescently labeled phalloidin was used to stain F-actin of muscles and visualize muscle patterns. Muscles are arranged in circular, longitudinal, and helicoidal orientations. Circular muscles were in the form of discreet rings around the pharynx and intestine, and contribute to the structure of the oral hood. Longitudinal muscles are largely concentrated on the ventral and ventrolateral sides of the body, where they aid in body flexion, including directional changes during ciliary swimming, body torsion, and escape responses. Helicoidal muscles, present as myocytes in left- and right-hand orientations, lie external of the circular bands and some of the longitudinal bands, and are hypothesized to counteract dilations of the pharynx and intestine during feeding. Extraordinary muscle orientations with undetermined functions include a pair of crossover muscles and a single semicircular muscle band at the caudal end. Accepted: 12 February 2001     

Locomotor Patterns in the Evolution of Actinopterygian Fishes

SYNOPSIS. Locomotor adaptations in actinopterygian fishes aredescribed for (a) caudal propulsion, used in cruising and sprintswimming, acceleration, and fast turns and (b) median and pairedfin propulsion used for slow swimming and in precise maneuver.Caudal swimming is subdivided into steady (time independent)and unsteady (time dependent acceleration and turning) locomotion. High power caudal propulsion is the major theme in actinopterygianswimming morphology because of its role in predator evasionand food capture. Non-caudal slow swimming appears to be secondaryand is not exploited before the Acanthopterygii. Optimal morphological requirements for unsteady swimming are(a) large caudal fin and general body area, (b) deep caudalpeduncle, often enhanced by posterior dorsal and anal fins,(c) an anterior stabilizing body mass and\or added mass, (d)flexible body and (e) large ratio of muscle mass to body mass.Optimal morphological requirements for steady swimming are (a)high aspect ratio caudal fin, (b) narrow caudal peduncle, (c)small total caudal area, (d) anterior stabilizing body massand added mass, and (e) a stiff body. Small changes in morphologycan have large effects on performance. Exclusive morphological requirements for steady versus unsteadyswimming are partially overcome using collapsible fins, butcompromises remain necessary. Morphologies favoring unsteadyperformance are a recurring theme in actinopterygian evolution.Successive radiations at chondrostean, halecostome and teleosteanlevels are associated with modifications in the axial and caudalskeleton. Strength of ossified structures probably limited maximum propulsionforces early in actinopterygian evolution, so that specializationsfor fast cruising (carangiform and thunmform modes) followedstructural advances especially in the caudal skeleton. No suchlimits apply to eel-like forms which consequently recur in successiveactinopterygian radiations. Slow swimming using mainly non-caudal propulsion probably firstoccurred among neopterygians in association with reduced andneutral buoyancy. Slow swimming adaptations can add to and extendthe scope of caudal swimming, but specialization is associatedwith reduced caudal swimming performance. Marked exploitationof slow swimming opportunities does not occur prior to the anterodorsallocation of pectoral and pelvic girdles and the vertical rotationof the base of the pectoral fin, as found in the Acanthopterygii.     

Center of mass motion in swimming fish: effects of speed and locomotor mode during undulatory propulsion

Studies of center of mass (COM) motion are fundamental to understanding the dynamics of animal movement, and have been carried out extensively for terrestrial and aerial locomotion. But despite a large amount of literature describing different body movement patterns in fishes, analyses of how the center of mass moves during undulatory propulsion are not available. These data would be valuable for understanding the dynamics of different body movement patterns and the effect of differing body shapes on locomotor force production. In the present study, we analyzed the magnitude and frequency components of COM motion in three dimensions (x: surge, y: sway, z: heave) in three fish species (eel, bluegill sunfish, and clown knifefish) swimming with four locomotor modes at three speeds using high-speed video, and used an image cross-correlation technique to estimate COM motion, thus enabling untethered and unrestrained locomotion. Anguilliform swimming by eels shows reduced COM surge oscillation magnitude relative to carangiform swimming, but not compared to knifefish using a gymnotiform locomotor style. Labriform swimming (bluegill at 0.5 body lengths/s) displays reduced COM sway oscillation relative to swimming in a carangiform style at higher speeds. Oscillation frequency of the COM in the surge direction occurs at twice the tail beat frequency for carangiform and anguilliform swimming, but at the same frequency as the tail beat for gymnotiform locomotion in clown knifefish. Scaling analysis of COM heave oscillation for terrestrial locomotion suggests that COM heave motion scales with positive allometry, and that fish have relatively low COM oscillations for their body size.     

Intersegmental coordination in invertebrates and vertebrates

How does the CNS coordinate muscle contractions between different body segments during normal locomotion? Work on several preparations has shown that this coordination relies on excitability gradients and on differences between ascending and descending intersegmental coupling. Abstract models involving chains of coupled oscillators have defined properties of coordinating circuits that would permit them to establish a constant intersegmental phase in the face of changing periods. Analyses that combine computational and experimental strategies have led to new insights into the cellular organization of intersegmental coordinating circuits and the neural control of swimming in lamprey, tadpole, crayfish and leech.     

The musculotendinous system of an anguilliform swimmer: Muscles, myosepta, dermis, and their interconnections in Anguilla rostrata

Eel locomotion is considered typical of the anguilliform swimming mode of elongate fishes and has received substantial attention from various perspectives such as swimming kinematics, hydrodynamics, muscle physiology, and computational modeling. In contrast to the extensive knowledge of swimming mechanics, there is limited knowledge of the internal body morphology, including the body components that contribute to this function. In this study, we conduct a morphological analysis of the collagenous connective tissue system, i.e., the myosepta and skin, and of the red muscle fibers that sustain steady swimming, focusing on the interconnections between these systems, such as the muscle-tendon and myosepta-skin connections. Our aim is twofold: (1) to identify the morphological features that distinguish this anguilliform swimmer from subcarangiform and carangiform swimmers, and (2) to reveal possible pathways of muscular force transmission by the connective tissue in eels. To detect gradual morphological changes along the trunk we investigated anterior (0.4L), midbody (0.6L), and posterior body positions (0.75L) using microdissections, histology, and three-dimensional reconstructions. We find that eel myosepta have a mediolaterally oriented tendon in each the epaxial and hypaxial regions (epineural or epipleural tendon) and two longitudinally oriented tendons (myorhabdoid and lateral). The latter two are relatively short (4.5-5% of body length) and remain uniform along a rostrocaudal gradient. The skin and its connections were additionally analyzed using scanning electron microscopy (SEM). The stratum compactum of the dermis consists of approximately 30 layers of highly ordered collagen fibers of alternating caudodorsal and caudoventral direction, with fiber angles of 60.51 +/- 7.05 degrees (n = 30) and 57.58 +/- 6.92 degrees (n = 30), respectively. Myosepta insert into the collagenous dermis via fiber bundles that pass through the loose connective tissue of the stratum spongiosum of the dermis and either weave into the layers of the stratum compactum (weaving fiber bundles) or traverse the stratum compactum (transverse fiber bundles). These fiber bundles are evenly distributed along the insertion line of the myoseptum. Red muscles insert into lateral and myorhabdoid myoseptal tendons but not into the horizontal septum or dermis. Thus, red muscle forces might be distributed along these tendons but will only be delivered indirectly into the dermis and horizontal septum. The myosepta-dermis connections, however, appear to be too slack for efficient force transmission and collagenous connections between the myosepta and the horizontal septum are at obtuse angles, a morphology that appears inadequate for efficient force transmission. Though the main modes of undulatory locomotion (anguilliform, subcarangiform, and carangiform) have recently been shown to be very similar with respect to their midline kinematics, we are able to distinguish two morphological classes with respect to the shape and tendon architecture of myosepta. Eels are similar to subcarangiform swimmers (e.g., trout) but are substantially different from carangiform swimmers (e.g., mackerel). This information, in addition to data from kinematic and hydrodynamic studies of swimming, shows that features other than midline kinematics (e.g., wake patterns, muscle activation patterns, and morphology) might be better for describing the different swimming modes of fishes.