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PubMed1995

Locomotion patterns in cerebral palsy syndromes.

Locomotion patterns were studied in 160 children with cerebral palsy. Ten patterns were distinguished, which were typical of the type of impairment: (1) crawling--mildly impaired children who will achieve independent walking; (2) creeping and crawling--diplegic children with moderate motor impairment; (3) creeping, never leading to independent walking--children with severe diplegia or tetraplegia; (4) bottom shuffling--children with hemiplegia and ataxia and minimal or mild motor impairment; (5) bunny-hopping--dyskinetic children with marked motor impairment but generally not mentally retarded; (6) rolling--severely diplegic and dyskinetic children; (7) other forms of locomotion, such as bridging or grub-type creeping, rare and typical of children with ataxic elements; (8) just walk--children with hemiplegia, diplegia, ataxic diplegia or ataxia, generally in cases of mild motor impairment but mental retardation; (9) just walk with aids--children with severe diplegia; and (10) no mobility. The locomotion pattern, age at onset and even manner of execution all influenced prognosis for walking. Severe deformity affected the choice of locomotion pattern. Though physiotherapy probably would not greatly influence the adoption of a particular locomotion pattern, early intervention might help prevent deformities.

Adolescent↗
PubMed1998

Cerebellar-induced locomotion: reticulospinal control of spinal rhythm generating mechanism in cats.

In a decerebrate cat (locomotor preparation), stimulation of a restricted region along the midline cerebellar white matter has been found to evoke generalized augmentation of postural muscle tone on a stationary surface (Asanome et al. 1998. Neurosci. Res. 30: 257-269) and "controlled" locomotion on the surface of a moving treadmill. Characteristics of cerebellar-evoked locomotion were similar to those of mesencephalic locomotor region-evoked "controlled" locomotion on the same animal. Microinjection of a neural tracer (CTb-HRP) into the lesioned stimulus site of the cerebellar white matter resulted in both retrograde labelling of cells in the fastigial nuclei, bilaterally, and anterograde labeling of fibers descending to the brain stem. These results indicated that the effective cerebellar stimulus site (cerebellar locomotor region) corresponded to the midline region of the hook bundle of Russell (Rasmussen, A. T., 1933. J. Comp. Neurol. 57: 165-197), through which crossed fastigioreticular, fastigiovestibular, and fastigiospinal fibers pass. In this study, contribution of reticulospinal systems to the control of cerebellar-evoked locomotion was extensively studied. By stimulating the cerebellar locomotor region and the MLR in the same animal, a majority of antidromically identified pontomedullary reticulospinal cells were synaptically activated. The results of the present study demonstrated that fastigial cells with crossed fastigioreticular fibers and reticulospinal fibers play a crucial role in the control of posture and locomotion in the locomotor preparation.

Animals↗
PubMedDate not supplied

Adaptive changes of locomotion after central and peripheral lesions.

This paper reviews findings on the adaptive changes of locomotion in cats after spinal cord or peripheral nerve lesions. From the results obtained after lesions of the ventral/ventrolateral pathways or the dorsal/dorsolateral pathways, we conclude that with extensive but partial spinal lesions, cats can regain voluntary quadrupedal locomotion on a treadmill. Although tract-specific deficits remain after such lesions, intact descending tracts can compensate for the lesioned tracts and access the spinal network to generate voluntary locomotion. Such neuroplasticity of locomotor control mechanisms is also demonstrated after peripheral nerve lesions in cats with intact or lesioned spinal cords. Some models have shown that recovery from such peripheral nerve lesions probably involves changes at the supra spinal and spinal levels. In the case of somesthesic denervation of the hindpaws, we demonstrated that cats with a complete spinal section need some cutaneous inputs to walk with a plantigrade locomotion, and that even in this spinal state, cats can adapt their locomotion to partial cutaneous denervation. Altogether, these results suggest that there is significant plasticity in spinal and supraspinal locomotor controls to justify the beneficial effects of early proactive and sustained locomotor training after central (Rossignol and Barbeau 1995; Barbeau et al. 1998) or peripheral lesions.

Adaptation, Physiological↗
PubMedDate not supplied

Functional interaction of dopamine and glutamate in the nucleus accumbens in the regulation of locomotion.

The interaction of dopamine and glutamate in the nucleus accumbens in the regulation of locomotion was investigated. Microinjection of N-methyl-D-aspartic acid (NMDA, a glutamatergic NMDA receptor agonist) or alpha-amino-3-hydroxy-5-methylisoxazole-4-propionate (AMPA, a quisqualic receptor agonist which is a glutamatergic non-NMDA receptor agonist) into the nucleus accumbens caused a substantial increase in locomotor activity. This increase in locomotor activity was significantly reduced by prior administration of the dopamine D2 agonist quinpirole, but not the D1 agonist, SKF 38393, into the same brain sites. The reduction in locomotion produced by quinpirole was dose dependent. Eight days after the ventral tegmental area was lesioned with 6-hydroxydopamine to destroy the dopamine projection and the axon terminals of the mesolimbic dopamine neurons in nucleus accumbens, the hyperkinetic effects produced by injections of NMDA and AMPA into the nucleus accumbens were substantially reduced. These results suggested that the glutamate agonist induced locomotion is mediated by dopamine. Thus, it appears that NMDA- or AMPA-induced locomotion is due to the activation of glutamate receptors on the mesolimbic dopamine terminals in the nucleus accumbens which release dopamine and subsequently increase locomotion.

Animals↗
PubMed1982

Energetics and mechanics of terrestrial locomotion.

This review addresses a simple question: How do muscles use the energy they consume during terrestrial locomotion? Using a comparative approach, it was found that the mass-specific rate of metabolic energy consumption changes by more than ten-fold with body size, while the mass-specific rate at which the muscles performed mechanical work did not change at all. It was also found that the rate of metabolic energy consumption increased linearly with speed, while the rate at which muscles performed mechanical work increased curvilinearly with speed (oc V1.53). We conclude from these observations that the rate at which animals consume metabolic energy during terrestrial locomotion is not determined by the rate at which their muscles perform mechanical work. Instead, the metabolic cost of generating muscular force over time (integral of F dt) appears to determine the metabolic cost of terrestrial locomotion. The cost of generating force increases with increasing speed and decreases with increasing body size in exactly the same manner as cost of locomotion. It is suggested that the metabolic cost of generating muscular force may be determined by the intrinsic velocity of shortening (i.e proportional to rates at which the cross-bridges between actin and myosin cycle) of the muscle motor units that are active during locomotion. Faster motor units are used both as animals increase speed and in equivalent muscles of smaller animals moving at the same speed. This suggestion is testable and future studies should determine whether or not it explains the higher costs of generating muscular force with increasing speed and decreasing body size.

Animals↗
PubMed2004

5-HT prolongs ventral root bursting via presynaptic inhibition of synaptic activity during fictive locomotion in lamprey.

Locomotor pattern generation is maintained by integration of the intrinsic properties of spinal central pattern generator (CPG) neurons in conjunction with synaptic activity of the neural network. In the lamprey, the spinal locomotor CPG is modulated by 5-HT. On a cellular level, 5-HT presynaptically inhibits synaptic transmission and postsynaptically inhibits a Ca2+-activated K+ current responsible for the slow afterhyperpolarization (sAHP) that follows action potentials in ventral horn neurons. To understand the contribution of these cellular mechanisms to the modulation of the spinal CPG, we have tested the effect of selective 5-HT analogues against fictive locomotion initiated by bath application of N-methyl-d-aspartate (NMDA). We found that the 5-HT1D agonist, L694-247, dramatically prolongs the frequency of ventral root bursting. Furthermore, we show that L694-247 presynaptically inhibits synaptic transmission without altering postsynaptic Ca2+-activated K+ currents. We also confirm that 5-HT inhibits synaptic transmission at concentrations that modulate locomotion. To examine the mechanism by which selective presynaptic inhibition modulates the frequency of fictive locomotion, we performed voltage- and current-clamp recordings of CPG neurons during locomotion. Our results show that 5-HT decreases glutamatergic synaptic drive within the locomotor CPG during fictive locomotion. Thus we conclude that presynaptic inhibition of neurotransmitter release contributes to 5-HT-mediated modulation of locomotor activity.

Animals↗
PubMed2005

Kinematic and EMG determinants in quadrupedal locomotion of a non-human primate (Rhesus).

We hypothesized that the activation patterns of flexor and extensor muscles and the resulting kinematics of the forelimbs and hindlimbs during locomotion in the Rhesus would have unique characteristics relative to other quadrupedal mammals. Adaptations of limb movements and in motor pool recruitment patterns in accommodating a range of treadmill speeds similar to other terrestrial animals in both the hindlimb and forelimb were observed. Flexor and extensor motor neurons from motor pools in the lumbar segments, however, were more highly coordinated than in the cervical segments. Unlike the lateral sequence characterizing subprimate quadrupedal locomotion, non-human primates use diagonal coordination between the hindlimbs and forelimbs, similar to that observed in humans between the legs and arms. Although there was a high level of coordination between hind- and forelimb locomotion kinematics, limb-specific neural control strategies were evident in the intersegmental coordination patterns and limb endpoint trajectories. Based on limb kinematics and muscle recruitment patterns, it appears that the hindlimbs, and notably the distal extremities, contribute more to body propulsion than the forelimbs. Furthermore, we found adaptive changes in the recruitment patterns of distal muscles in the hind- and forelimb with increased treadmill speed that likely correlate with the anatomical and functional evolution of hand and foot digits in monkeys. Changes in the properties of both the spinal and supraspinal circuitry related to stepping, probably account for the peculiarities in the kinematic and EMG properties during non-human primate locomotion. We suggest that such adaptive changes may have facilitated evolution toward bipedal locomotion.

Analysis of Variance↗
PubMed1978

Controlled locomotion in the mesencephalic cat: distribution of facilitatory and inhibitory regions within pontine tegmentum.

1. The contribution of postural tonus to controlled locomotion elicited by the stimulation of mesencephalic locomotor region (MLR) was studied in the acute precolicular-postmammillary decerebrate (mesencephalic) cat. 2. A microelectrode was placed in the unilateral MLR and another was placed systematically at 1-mm increments throughout the pons (H--4 to H--1O) at level ranging from P2 to P11 dorsoventrally and mediolaterally from 0 to L or R6. Depending on the general condition of the animal, stimuli through this second electrode were delivered preceding, succeeding, or simultaneous with the MLR stimulation. 3. Stimulation of the ventral part of the caudal tegmental field (P3 to P9, H--6 to H--8) increased extensor tonus of the hindlimbs, as assessed by recording muscle activity. Concomitant stimulation of this region converted MLR-elicited hindlimb stepping to coordinated four-legged locomotion and also elicited locomotion even when stimulation of the MLR alone failed to elicit locomotion. Stimulation of this ventral tegmental region alone at a larger stimulus intensity elicited spastic locomotor movements associated with a substantial increase in extensor tonus. 4. Stimulation of the lateral tegmented field surrounding the pontine locomotor region (PLR) also facilitated MLR effects, but had a relatively weaker facilitatory effect on postural tonus then stimulation of the ventrocaudal tegmental field. PLR stimulation alone was also ineffective when postural tonus was not well maintained. 5 Stimulation of the dorsal part of caudal tegmental field (P3 to P9, H--4 to H--6) in its midline dramatically decreased extensor tonus of the hindlimbs. MLR-elicited controlled locomotion was completely suppressed during concomitant stimulation of this inhibitory region. 6. These results indicated clearly that the degree of existing postural tonus greatly affects MLR-elicited locomotor movements and that an increase in postural tonus and an activation of the spinal stepping generator are not separate phenomenia.

Animals↗
PubMed1984

Synaptic transmission from muscle afferents during fictive locomotion in the mesencephalic cat.

Modulation of synaptic potentials produced by electrical stimulation of low-threshold muscle afferents in lumbar alpha-motoneurons innervating knee and ankle muscles was studied by intracellular recording during "fictive locomotion" induced by stimulating the mesencephalic locomotor region (MLR) in paralyzed, mesencephalic cats. Averaging postsynaptic potentials in different phases of the fictive step cycle indicated that relatively little amplitude modulation occurred. In nearly half of the 38 motoneurons analyzed, there was a statistically significant tendency for excitatory postsynaptic potentials (EPSPs) to increase in amplitude during the depolarized phase of the oscillation in the membrane potential produced during fictive locomotion (locomotor-drive potential). In 8% the EPSPs decreased under the same conditions, while the rest displayed a constant amplitude during all phases of the fictive step cycle. Only three cells showed a distinct second peak in the EPSP at a latency consistent with transmission in a di- or trisynaptic pathway. Late inhibitory postsynaptic potentials (IPSPs) were also rarely observed. Thus oligosynaptic pathways from muscle afferents to the motoneuron groups studied are not prominent during the locomotor cycle in this preparation. We suggest that the marked modulation of monosynaptic reflex amplitude observed in mesencephalic cats (1) arises mainly from the effects of the locomotor-drive potential in bringing the cells closer to threshold during some phases of locomotion. Specific modulation during fictive locomotion of transmission in pathways from muscle afferents, which has been demonstrated for cutaneous pathways (28), was not observed. The implications of these results for the control of locomotion are discussed briefly.

Animals↗
PubMed2000

Maintenance of rat head direction cell firing during locomotion in the vertical plane.

Previous studies have identified a subset of neurons in the rat anterodorsal thalamus (ADN) that encode head direction (HD) in absolute space and may be involved in navigation. These HD cells discharge selectively when the rat points its head in a specific direction (the preferred firing direction) in the horizontal plane. HD cells are typically recorded during free movement about a single horizontal surface. The current experiment examined how HD cell firing was influenced by 1) locomotion in the vertical plane and 2) locomotion on two different horizontal surfaces separated in height. Rats were trained in a cylindrical enclosure containing a single polarizing cue card attached to the cylinder wall, covering approximately 100 degrees of arc. The enclosure contained two horizontal surfaces: the cylinder floor and an annulus around the cylinder top 76 cm above the floor. A 90 degrees vertical mesh ladder that could be affixed at any angular position on the cylinder wall allowed the rats to locomote back and forth between the two horizontal surfaces. Rats were trained to retrieve food pellets on the cylinder floor as well as climb the mesh ladder to retrieve food pellets on the annulus. HD cell activity was monitored as the rat traversed the horizontal and vertical surfaces of the apparatus. When the angular position of the mesh corresponded to the cell's preferred firing direction, the HD cells maintained their peak discharge rate as the rat climbed up the mesh, but did not fire when the rat climbed down the mesh. In contrast, when the mesh was positioned 180 degrees opposite the preferred firing direction, HD cells did not fire when the rat climbed up the mesh, but exhibited maximal firing when the rat climbed down the mesh. When the mesh was placed 90 or 270 degrees from the preferred firing direction, HD cells exhibited background firing rates during climbing up or down the mesh. While preferred firing directions were maintained between the two horizontal surfaces, peak firing rate increased significantly (approximately 30%) on the annulus as compared with the cylinder floor. These data demonstrate that HD cells continue to discharge in the vertical plane if the vertical locomotion began with the rat's orientation corresponding to the preferred firing direction. One model consistent with these data are that HD cells define the horizontal reference frame as the animal's plane of locomotion. Further, we propose that HD cell firing, as viewed within a three-dimensional coordinate system, can be characterized as the surface of a hemitorus.

Animals↗
PubMed2002

Intracellular QX-314 causes depression of membrane potential oscillations in lamprey spinal neurons during fictive locomotion.

Spinal neurons undergo large cyclic membrane potential oscillations during fictive locomotion in lamprey. It was investigated whether these oscillations were due only to synaptically driven excitatory and inhibitory potentials or if voltage-dependent inward conductances also contribute to the depolarizing phase by using N-(2,6-dimethylphenyl carbamoylmethyl)triethylammonium bromide (QX-314) administered intracellularly during fictive locomotion. QX-314 intracellularly blocks inactivating and persistent Na+ channels, and in some neurons, effects on certain other types of channels have been reported. To detail the effects of QX-314 on Na+ and Ca2+ channels, we used dissociated lamprey neurons recorded under whole cell voltage clamp. At low intracellular concentrations of QX-314 (0.2 mM), inactivating Na+ channels were blocked and no effects were exerted on Ca2+ channels (also at 0.5 mM). At 10 mM QX-314, there was, however a marked reduction of I(Ca). In the isolated spinal cord of the lamprey, fictive locomotion was induced by superfusing the spinal cord with Ringer's solution containing N-methyl-D-aspartate (NMDA), while recording the locomotor activity from the ventral roots. Simultaneously, identified spinal neurons were recorded intracellularly, while infusing QX-314 from the microelectrode. Patch electrodes cannot be used in the intact spinal cord, and therefore "sharp" electrodes were used. The amplitude of the oscillations was consistently reduced by 20-25% in motoneurons (P < 0.05) and unidentified spinal neurons (P < 0.005). The onset of the effect started a few minutes after impalement and reached a stable level within 30 min. These effects thus show that QX-314 causes a reduction in the amplitude of membrane potential oscillations during fictive locomotion. We also investigated whether QX-314 could affect glutamate currents by applying short pulses of glutamate from an extracellular pipette. No changes were observed. We also found no evidence for a persistent Na+ current in dissociated neurons, but these cells have a much-reduced dendritic tree. The results indicate that there is an inward conductance, which is sensitive to QX-314, during membrane potential oscillations that "boosts" the synaptic drive during fictive locomotion. Taken together, the results suggest that inactivating Na+ channels contribute to this inward conductance although persistent Na+ channels, if present on dendrites, could possibly also contribute to shaping the membrane potential oscillations.

Action Potentials↗
PubMed2006

Dynamic sensorimotor interactions in locomotion.

Locomotion results from intricate dynamic interactions between a central program and feedback mechanisms. The central program relies fundamentally on a genetically determined spinal circuitry (central pattern generator) capable of generating the basic locomotor pattern and on various descending pathways that can trigger, stop, and steer locomotion. The feedback originates from muscles and skin afferents as well as from special senses (vision, audition, vestibular) and dynamically adapts the locomotor pattern to the requirements of the environment. The dynamic interactions are ensured by modulating transmission in locomotor pathways in a state- and phase-dependent manner. For instance, proprioceptive inputs from extensors can, during stance, adjust the timing and amplitude of muscle activities of the limbs to the speed of locomotion but be silenced during the opposite phase of the cycle. Similarly, skin afferents participate predominantly in the correction of limb and foot placement during stance on uneven terrain, but skin stimuli can evoke different types of responses depending on when they occur within the step cycle. Similarly, stimulation of descending pathways may affect the locomotor pattern in only certain phases of the step cycle. Section ii reviews dynamic sensorimotor interactions mainly through spinal pathways. Section iii describes how similar sensory inputs from the spinal or supraspinal levels can modify locomotion through descending pathways. The sensorimotor interactions occur obviously at several levels of the nervous system. Section iv summarizes presynaptic, interneuronal, and motoneuronal mechanisms that are common at these various levels. Together these mechanisms contribute to the continuous dynamic adjustment of sensorimotor interactions, ensuring that the central program and feedback mechanisms are congruous during locomotion.

Animals↗
PubMed1991

Central command increases sympathetic nerve activity during spontaneous locomotion in cats.

A controversial issue in exercise physiology is the relative contribution of central command versus afferent input from contracting muscles and baroreceptors in the regulation of sympathetic nerve activity (SNA) during exercise. Recent studies of exercising humans have suggested that central command increases cutaneous sympathetic sudomotor nerve activity but have challenged the concept that central command contributes importantly to increases in sympathetic vasoconstrictor nerve activity to skin and skeletal muscle. The purpose of this study was to examine the influence of central command on renal SNA and lumbar SNA during spontaneous locomotion in decorticate cats. Unanesthetized decorticate cats that developed locomotion spontaneously or during electrical stimulation of the subthalamic locomotor region were studied in the presence and absence of input from skeletal muscle and baroreceptor afferents. Spontaneous rhythmic locomotion in the unparalyzed state was associated with significant increases in mean arterial pressure (MAP) from 106 +/- 10 to 133 +/- 11 mm Hg (p less than 0.05) and increases in renal SNA of 301 +/- 100% (p less than 0.05). During spontaneous fictive rhythmic locomotion in paralyzed cats, there were also significant (p less than 0.05) increases in MAP (43 +/- 6%), renal SNA (183 +/- 32%), and lumbar SNA (223 +/- 83%). Baroreceptor denervation did not attenuate increases in MAP, renal SNA, and lumbar SNA during locomotion. During electrical stimulation of the subthalamic locomotor region in paralyzed cats, MAP increased by 43 +/- 17% (p less than 0.05), and renal SNA increased by 175 +/- 47% (p less than 0.05). These findings indicate that central command is capable of increasing sympathetic neural drive in unanesthetized decorticate cats. This increase in sympathetic drive occurs even in the absence of feedback from contracting muscles or from arterial and cardiopulmonary baroreceptors.

Animals↗
PubMed2005

Hindlimb function in the alligator: integrating movements, motor patterns, ground reaction forces and bone strain of terrestrial locomotion.

Alligator hindlimbs show high torsional loads during terrestrial locomotion, in sharp contrast to the bending or axial compressive loads that predominate in animals that use parasagittal limb movements. The present study integrates new data on hindlimb muscle function with previously obtained data on hindlimb kinematics, motor patterns, ground reaction forces and bone strain in order to (1) assess mechanisms underlying limb bone torsion during non-parasagittal locomotion in alligators and (2) improve understanding of hindlimb dynamics during terrestrial locomotion. Three dynamic stance phase periods were recognized: limb-loading, support-and-propulsion, and limb-unloading phases. Shear stresses due to torsion were maximized during the limb-loading phase, during which the ground reaction force (GRF) and caudofemoralis (CFL) muscles generated opposing moments about the femur. Hindlimb retraction during the subsequent stance-and-propulsion phase involves substantial medial rotation of the femur, powered largely by coordinated action of the GRF and CFL. Several muscles that actively shorten to flex and extend limb joints during stance phase in sprawling and erect quadrupeds act in isometric or even eccentric contraction in alligators, stabilizing the knee and ankle during the support-and-propulsion phase. Motor patterns in alligators reveal the presence of local and temporal segregation of muscle functions during locomotion with muscles that lie side by side dedicated to performing different functions and only one of 16 muscles showing clear bursts of activity during both stance and swing phases. Data from alligators add to other recent discoveries that homologous muscles across quadrupeds often do not move joints the same way as is commonly assumed. Although alligators are commonly considered models for early semi-erect tetrapod locomotion, many aspects of hindlimb kinematics, muscle activity patterns, and femoral loading patterns in alligators appear to be derived in alligators rather than reflecting an ancestral semi-erect condition.

Alligators and Crocodiles↗
PubMed1988

Muscular mechanisms of snake locomotion: an electromyographic study of the sidewinding and concertina modes of Crotalus cerastes, Nerodia fasciata and Elaphe obsoleta.

Synchronized electromyography and cinematography were used to determine the muscle activity of colubroid snakes during sidewinding and concertina locomotion. The primary muscles studied were the three largest, most superficial epaxial muscles: the Mm. semispinalis-spinalis, longissimus dorsi and iliocostalis. Sidewinding locomotion of Nerodia fasciata and Crotalus cerastes was the result of continuous posterior propagation of contractile blocks consisting of several adjacent muscle segments. During sidewinding, the activity of the M. longissimus dorsi and M. iliocostalis was primarily unilateral, beginning when a body region was convex and ending when it was maximally concave on the side of the active muscle. Unilateral activity of the M. semispinalis-spinalis correlated with lateral flexion in addition to bilateral activity that correlated with dorsiflexion of the vertebral column. During concertina locomotion of N. fasciata and Elaphe obsoleta, muscle activity also involved blocks of several simultaneously active adjacent muscle segments, but all major activity was unilateral and was not propagated posteriorly in a simple continuous fashion. Muscle activity during concertina locomotion correlated either with lateral flexion towards the side of the active muscle or with the maintenance of static contact with the sides of a tunnel. The number of simultaneously active adjacent muscle segments and the maximum duration of continuous muscle activity varied significantly between Nerodia and Elaphe and among the different widths of tunnels. Theoretical considerations combined with observed differences suggest that the more elongate body of Elaphe is advantageous for performing concertina locomotion. There was no consistent evidence that nonhomologous muscles with tendinous interconnections functioned as single units during either of these two locomotor modes. Although individual segments of the studied epaxial muscles span several vertebrae, via long, tendinous connections, consistent kinematic correlations with muscle activity were observed only between the contractile portion of a muscle segment and the vertebrae adjacent to that contractile portion.

Animals↗
PubMed1990

The work and energetic cost of locomotion. I. The effects of limb mass distribution in quadrupeds.

Does limb design influence the cost of locomotion in quadrupedal mammals? If not, morphologists must dismiss the economy of locomotion from consideration when assessing the adaptive factors shaping limb structure. Several studies have recently used externally applied loads to demonstrate a relationship between limb mass distribution and energy costs in human subjects. It is not clear whether a similar correlation would hold for quadrupeds, given their very different gaits. The present study addresses this question by measuring the rate of oxygen consumption in domestic dogs running on a treadmill with mass added either to the limbs or to the back. Trials with no additional mass were used as a control. The use of externally applied loads has the advantage of allowing limb mass to be altered in a system in which other aspects of physiology that might influence cost of locomotion are held constant. The cost of adding mass to the limbs in dogs was found to be significantly greater than that of adding it at the center of mass. Limb mass distribution does affect the cost of locomotion in quadrupeds. A comparison of the results from a variety of studies in which the energetic cost of adding external loads has been measured in animals across a wide size range suggests a qualitative difference in the factors determining the cost of locomotion in large and small animals.

Animals↗
PubMed2000

Patterns of white muscle activity during terrestrial locomotion in the American eel (Anguilla rostrata).

Eels (Anguilla rostrata) are known to make occasional transitory excursions into the terrestrial environment. While on land, their locomotor kinematics deviate drastically from that observed during swimming. In this study, electromyographic (EMG) recordings were made from white muscle at various longitudinal positions in eels performing undulatory locomotion on land to determine the muscle activity patterns underlying these terrestrial movements. As during swimming, eels propagate a wave of muscle activity from anterior to posterior during terrestrial locomotion. However, the intensity of EMG bursts is much greater on land (on average approximately five times greater than in water). In addition, anteriorly located musculature has higher-intensity EMG bursts than posteriorly located muscle during locomotion on land. EMG duty cycle (burst duration relative to undulatory cycle time) is significantly affected by longitudinal position during terrestrial locomotion, and duty cycles are significantly greater on land (0.4-0.5 cycles) than in water (0. 2-0.3 cycles). Finally, as in swimming, a phase shift in the timing of muscle activity exists such that posteriorly located muscle fibers become activated earlier in their strain cycle than do more anteriorly located fibers. However, fibers become activated much later in their muscle strain cycle on land than in water. Therefore, it is clear that, while eels propagate a wave of muscle activity posteriorly to generate backward-traveling waves that generate propulsive thrust both in water and on land, the specific patterns of timing and the intensity of muscle activity are substantially altered depending upon the environment. This suggests that physical differences in an animal's external environment can play a substantial role in affecting the motor control of locomotion, even when similar structures are used to generate the propulsive forces.

Anguilla↗
PubMed2001

Pectoral fin locomotion in batoid fishes: undulation versus oscillation.

This study explores the dichotomy between undulatory (passing multiple waves down the fin or body) and oscillatory (flapping) locomotion by comparing the kinematics of pectoral fin locomotion in eight species of batoids (Dasyatis americana, D. sabina, D. say, D. violacea, Gymnura micrura, Raja eglanteria, Rhinobatos lentiginosus and Rhinoptera bonasus) that differ in their swimming behavior, phylogenetic position and lifestyle. The goals of this study are to describe and compare the pectoral fin locomotor behavior of the eight batoid species, to clarify how fin movements change with swimming speed for each species and to analyze critically the undulation/oscillation continuum proposed by Breder using batoids as an example. Kinematic data were recorded for each species over a range of swimming velocities (1-3 disc lengths s(-1)). The eight species in this study vary greatly in their swimming modes. Rhinobatos lentiginosus uses a combination of axial-based and pectoral-fin-based undulation to move forward through the water, with primary thrust generated by the tail. The pectoral fins are activated in short undulatory bursts for increasing swimming speed and for maneuvering. Raja eglanteria uses a combination of pectoral and pelvic locomotion, although only pectoral locomotion is analyzed here. The other six species use pectoral locomotion exclusively to propel themselves through the water. Dasyatis sabina and D. say have the most undulatory fins with an average of 1.3 waves per fin length, whereas Rhinoptera bonasus has the most oscillatory fin behavior with 0.4 waves per fin length. The remaining species range between these two extremes in the degree of undulation present on their fins. There is an apparent trade-off between fin-beat frequency and amplitude. Rhinoptera bonasus has the lowest frequency and the highest fin amplitude, whereas Rhinobatos lentiginosus has the highest frequency and the lowest amplitude among the eight species examined. The kinematic variables that batoids modify to change swimming velocity vary among different species. Rhinobatos lentiginosus increases its tail-beat frequency to increase swimming speed. In contrast, the four Dasyatis species increase swimming speed by increasing frequency and wavespeed, although D. americana also changes wave number. Raja eglanteria modifies its swimming velocity by changing wavespeed and wave number. Rhinoptera bonasus increases wavespeed, Gymnura micrura decreases wave number, and both Rhinoptera bonasus and Gymnura micrura increase fin-tip velocity to increase swimming velocity. Batoid species fall onto a continuum between undulation and oscillation on the basis of the number of waves present on the fins.

Animals↗