Locomotor Cycle Research Articles

A modelling study of locomotion-induced hyperpolarization of voltage threshold in cat lumbar motoneurones.

During fictive locomotion the excitability of adult cat lumbar motoneurones is increased by a reduction (a mean hyperpolarization of approximately 6.0 mV) of voltage threshold (Vth) for action potential (AP) initiation that is accompanied by only small changes in AP height and width. Further examination of the experimental data in the present study confirms that Vth lowering is present to a similar degree in both the hyperpolarized and depolarized portions of the locomotor step cycle. This indicates that Vth reduction is a modulation of motoneurone membrane currents throughout the locomotor state rather than being related to the phasic synaptic input within the locomotor cycle. Potential ionic mechanisms of this locomotor-state-dependent increase in excitability were examined using three five-compartment models of the motoneurone innervating slow, fast fatigue resistant and fast fatigable muscle fibres. Passive and active membrane conductances were set to produce input resistance, rheobase, afterhyperpolarization (AHP) and membrane time constant values similar to those measured in adult cat motoneurones in non-locomoting conditions. The parameters of 10 membrane conductances were then individually altered in an attempt to replicate the hyperpolarization of Vth that occurs in decerebrate cats during fictive locomotion. The goal was to find conductance changes that could produce a greater than 3 mV hyperpolarization of Vth with only small changes in AP height (< 3 mV) and width (< 1.2 ms). Vth reduction without large changes in AP shape could be produced either by increasing fast sodium current or by reducing delayed rectifier potassium current. The most effective Vth reductions were achieved by either increasing the conductance of fast sodium channels or by hyperpolarizing the voltage dependency of their activation. These changes were particularly effective when localized to the initial segment. Reducing the conductance of delayed rectifier channels or depolarizing their activation produced similar but smaller changes in Vth. Changes in current underlying the AHP, the persistent Na(+) current, three Ca(2+) currents, the "h" mixed cation current, the "A" potassium current and the leak current were either ineffective in reducing Vth or also produced gross changes in the AP. It is suggested that the increased excitability of motoneurones during locomotion could be readily accomplished by hyperpolarizing the voltage dependency of fast sodium channels in the axon hillock by a hitherto unknown neuromodulatory action.

Hypaxial muscle activity during running and breathing in dogs.

The axial muscles of terrestrial vertebrates serve two potentially conflicting functions, locomotion and lung ventilation. To differentiate the locomotor and ventilatory functions of the hypaxial muscles in mammals, we examined the locomotor and ventilatory activity of the trunk muscles of trotting dogs under two conditions: when the ventilatory cycle and the locomotor cycle were coupled and when they were uncoupled. Patterns of muscle-activity entrainment with locomotor and ventilatory events revealed (i) that the internal and external abdominal oblique muscles performed primarily locomotor functions during running yet their activity was entrained to expiration when the dogs were standing, (ii) that the internal and external intercostal, external oblique thoracic and transversus abdominis muscles performed both locomotor and respiratory functions simultaneously, (iii) that the parasternal internal intercostal muscle performed a primarily respiratory function (inspiration) and (iv) that the deep pectoralis and longissimus dorsi muscles performed only locomotor functions and were not active while the dogs were standing still. We conclude that the dual function of many hypaxial muscles may produce functional conflicts during running. The redundancy and complexity of the respiratory musculature as well as the particular pattern of respiratory-locomotor coupling in quadrupedal mammals may circumvent these conflicts or minimize their impact on respiration.

Selectivity of the central control of sensory information in the mammalian spinal cord.

Afferent feedback from muscle proprioceptors, as well as movement-induced activation of skin receptors plays an important role in the patterning of motor activity for stepping and postural control. An important component in this control is the presynaptic GABAergic modulation of the synaptic effectiveness of muscle and cutaneous afferents, known to change in phase with the locomotor cycle, during the execution of voluntary movements, or after a peripheral nerve injury. Recent electrophysiological studies, together with ultrastructural observations, indicate that the distribution of GABAa synapses in the intraspinal arborizations of muscle spindle and tendon organ afferents is not homogeneous. Namely, that some collaterals are the targets of one, or more, GABAergic interneurones, while other collaterals of the same fibre receive no GABAergic connections. In addition, both PAD and inhibition of PAD have a local character. This allows, at least in principle, decoupling the information arising from common sensory inputs. A spatially restricted modulation of PAD could play a significant role in the adjustment of the synaptic effectiveness of Ia afferents at the onset of voluntary contractions in humans, during movement-induced stimulation of the skin, or during the compensation of motor activity following partial denervation of muscles. Changes in the synchronization of the PAD-mediating interneurones can also have a profound effect on the information transmitted by a given set of afferent fibres. Data are presented that in the anesthetized cat, variation in the spontaneous activity of a population of dorsal horn neurones in laminae III-VI, that respond to stimulation of low-threshold cutaneous afferents, produce correlated fluctuations of monosynaptic reflexes by means of pre- and postsynaptic mechanisms. It is suggested that correlated changes in the level of PAD can also play a significant role in the presynaptic adjustment of the synaptic effectiveness of the afferent fibres during specific motor tasks.

Central pattern generation of locomotion: a review of the evidence.

Neural networks in the spinal cord, referred to as "central pattern generators" (CPGs), are capable of producing rhythmic movements, such as swimming, walking, and hopping, even when isolated from the brain and sensory inputs. This article reviews the evidence for CPGs governing locomotion and addresses other factors, including supraspinal, sensory, and neuromodulatory influences, that interact with CPGs to shape the final motor output. Supraspinal inputs play a major role not only in initiating locomotion but also in adapting the locomotor pattern to environmental and motivational conditions. Sensory afferents involved in muscle and cutaneous reflexes have important regulatory functions in preserving balance and ensuring stable phase transitions in the locomotor cycle. Neuromodulators evoke changes in cellular and synaptic properties of CPG neurons, conferring flexibility to CPG circuits. Briefly addressed is the interaction of CPG networks to produce intersegmental coordination for locomotion. Evidence for CPGs in humans is reviewed, and although a comprehensive clinical review is not an objective, examples are provided of animal and human studies that apply knowledge of CPG mechanisms to improve locomotion. The final section deals with future directions in CPG research.

Epaxial muscle function in trotting dogs.

One of the features that distinguish mammals from other groups of terrestrial vertebrates is the structure and relative size of their epaxial muscles. Yet we have only a superficial understanding of the role these muscles play in locomotion. To address their locomotor function, we recorded the electrical activity of the iliocostalis, longissimus dorsi and multifidus muscles of trotting dogs. Activity was monitored at both lumbar and thoracic sites. To develop and evaluate hypotheses of epaxial muscle function, we quantified footfall patterns and sagittal trunk kinematics from high-speed videos, and the magnitude and orientation of ground reaction forces from force-plate recordings. All three epaxial muscles tended to exhibit a double-bursting (biphasic) activity pattern, with the exception of the iliocostalis muscle at the thoracic site (which was uniphasic). In general, a large burst of activity in each muscle occurred during the second half of the support phase of the ipsilateral hindlimb, and was active for an average of 30% of the locomotor cycle. A smaller burst of activity occurred during the second half of the support phase of the contralateral hindlimb, and was active for an average of 15% of the locomotor cycle. Analysis of ground reaction forces and sagittal trunk kinematics led us to the hypothesis that the epaxial muscles do not directly stabilize the trunk against the vertical and horizontal components of the ground reaction force. Instead, the epaxial muscles appear to counteract the tendency of the trunk to rebound (flex) in the sagittal plane during the latter half of the support phase. This hypothesis of epaxial muscle function was supported by loading experiments performed on the longissimus dorsi muscle in the lumbar region.

Distinctive patterns of static and dynamic gamma motor activity during locomotion in the decerebrate cat.

Simultaneous recordings were made from gamma (gamma) motor axons and from muscle spindle afferents of the medial gastrocnemius (MG) muscle during locomotion in decerebrate cats. The gamma-neurons were identified as static or dynamic (gammas or gammad) by correlating their behaviour during midbrain stimulation with changes in muscle spindle afferent responses to muscle stretch. On the basis of their behaviour during locomotion, gammas neurons could be divided into two groups. One group (type-1) showed strongly and smoothly modulated discharge increasing in parallel with the active muscle shortening in ankle extension, but with phase advance. The other group (type-2) also showed a modulated pattern, but with increased firing centred on the flexion phase. The proportions of the two were 13 type-1 and 7 type-2. The type-1 firing pattern accurately predicted the difference in firing frequency for secondary afferents obtained by subtracting from the recordings made during active movements the response of the same units to the movements repeated passively in the absence of fusimotor activity. The type-2 pattern also became consistent with the difference signal, when operated on by a phase lag appropriate to the effects of bag2 intrafusal fibres. These results suggest that there may be some degree of separate control of chain and bag2 intrafusal fibres. The discharge of gammad axons was also found to fluctuate with the locomotor cycle, with a pattern very distinct from that of the gammas records. The gammad firing frequency rose very suddenly from zero to a maximum at the onset of muscle shortening and continued into the beginning of lengthening. The term 'interrupted' discharge is suggested as a useful description. The timing of this discharge was shown to be appropriate for sensitising the primary afferents to detect the onset of stretch.

The mechanics of locomotion in the squid Loligo pealei: locomotory function and unsteady hydrodynamics of the jet and intramantle pressure.

High-speed, high-resolution digital video recordings of swimming squid (Loligo pealei) were acquired. These recordings were used to determine very accurate swimming kinematics, body deformations and mantle cavity volume. The time-varying squid profile was digitized automatically from the acquired swimming sequences. Mantle cavity volume flow rates were determined under the assumption of axisymmetry and the condition of incompressibility. The data were then used to calculate jet velocity, jet thrust and intramantle pressure, including unsteady effects. Because of the accurate measurements of volume flow rate, the standard use of estimated discharge coefficients was avoided. Equations for jet and whole-cycle propulsive efficiency were developed, including a general equation incorporating unsteady effects. Squid were observed to eject up to 94 % of their intramantle working fluid at relatively high swimming speeds. As a result, the standard use of the so-called large-reservoir approximation in the determination of intramantle pressure by the Bernoulli equation leads to significant errors in calculating intramantle pressure from jet velocity and vice versa. The failure of this approximation in squid locomotion also implies that pressure variation throughout the mantle cannot be ignored. In addition, the unsteady terms of the Bernoulli equation and the momentum equation proved to be significant to the determination of intramantle pressure and jet thrust. Equations of propulsive efficiency derived for squid did not resemble Froude efficiency. Instead, they resembled the equation of rocket motor propulsive efficiency. The Froude equation was found to underestimate the propulsive efficiency of the jet period of the squid locomotory cycle and to overestimate whole-cycle propulsive efficiency when compared with efficiencies calculated from equations derived with the squid locomotory apparatus in mind. The equations for squid propulsive efficiency reveal that the refill period of squid plays a greater role, and the jet period a lesser role, in the low whole-cycle efficiencies predicted in squid and similar jet-propelled organisms. These findings offer new perspectives on locomotory hydrodynamics, intramantle pressure measurements and functional morphology with regard to squid and other jet-propelled organisms.

The role of putative glutamatergic neurons and their connections in the locomotor central pattern generator of the mollusk, Clione limacina

In the pteropod mollusk Clione limacina, locomotor rhythm is produced by the central pattern generator (CPG), due mainly to the activity of interneurons of groups 7 (active in the phase of the dorsal flexion of the wings) and 8 (active in the phase of the ventral flexion). Each of these groups excites the neurons active in the same phase of the locomotor cycle, and inhibits the neurons of the opposite phase. In this work, the nature of connections formed by group 7 interneurons was studied. Riluzole (2-amino-6-trifluoro-methoxybenzothiazole), which is known to inhibit the presynaptic release of glutamate, suppressed the action of the type 7 interneurons onto the follower neurons of the same and of the antagonistic phase of the locomotor cycle. The main pattern of rhythmic activity of CPG with alternation of two phases could be maintained after suppression of inhibitory connections from group 7 interneurons to antagonistic neurons. This suggests redundancy of the mechanisms controlling swimming rhythm generation, which ensures the reliable operation of the system.

Spike coding during locomotor network activity in ventrally located neurons in the isolated spinal cord from neonatal rat.

To characterize spike coding in spinal neurons during rhythmic locomotor activity, we recorded from individual cells in the lumbar spinal cord of neonatal rats by using the on-cell patch-clamp technique. Locomotor activity was induced by N-methyl-D aspartate (NMDA) and 5-hydroxytryptamine (5-HT) and monitored by ventral root recording. We made an estimator based on the assumption that the number of spikes arriving during two halves of the locomotor cycle could be a code used by the neuronal network to distinguish between the halves. This estimator, termed the spike contrast, was calculated as the difference between the number of spikes in the most and least active half of an average cycle. The root activity defined the individual cycles and the positions of the spikes were calculated relative to these cycles. By comparing the average spike contrast to the spike contrast in noncyclic, randomized spike trains we found that approximately one half the cells (19 of 42) contained a significant spike contrast, averaging 1.25 +/- 0.23 (SE) spikes/cycle. The distribution of spike contrasts in the total population of cells was exponential, showing that weak modulation was more typical than strong modulation. To investigate if this low spike contrast was misleading because a higher spike contrast averaged out by occurring at different positions in the individual cycles we compared the spike contrast obtained from the average cycle to its maximal value in the individual cycles. The value was larger (3.13 +/- 0.25 spikes) than the spike contrast in the average cycle but not larger than the spike contrast in the individual cycles of a random, noncyclic spike trains (3.21 +/- 0.21 spikes). This result suggested that the important distinction between cyclic and noncyclic cells was only the repeated cycle position of the spike contrast and not its magnitude. Low spike frequencies (5.2 +/- 0.82 spikes/cycle, that were on average 3.5 s long) and a minimal spike interval of 100-200 ms limited the spike contrast. The standard deviation (SD) of the spike contrast in the individual neurons was similar to the average spike contrasts and was probably stochastic because the SDs of the simulated, noncyclic spike trains were also similar. In conclusion we find a highly distributed and variable locomotor related cyclic signal that is represented in the individual neurons by very few spikes and that becomes significant only because the spike contrast is repeated at a preferred phase of the locomotor cycle.

Population Reconstruction of the Locomotor Cycle From Interneuron Activity in the Mammalian Spinal Cord

Lesion studies have shown that neuronal networks in the ventromedial regions of the neonatal rat spinal cord are critical for the production of locomotion. We examined whether the locomotor cycle could be accurately predicted based on the activity recorded in a population of spinal interneurons located in these regions during pharmacologically induced locomotion. We used a Bayesian probabilistic reconstruction procedure to predict the most likely phase of locomotion given the observed activity in the neuronal population. The population reconstruction was able to predict the correct locomotor phase with high accuracy using a relatively small number of neurons. This result demonstrates that although the spike activity of individual spinal interneurons in the ventromedial region is weak and varies from cycle to cycle, the locomotor phase can be accurately predicted when information from the population is combined. This result is consistent with the proposed involvement of interneurons within these regions of the spinal cord in the production of locomotion.

The Integration of Ventilation and Locomotion in Archosaurs

Movements of the pelvis have recently been found to contribute to ventilation in both crocodilians and birds. Alligators have a kinetic pelvis in which the ischiopubic and ischiotruncus muscles rotate the pubic bones ventrally to increase abdominal volume and thereby facilitate inspiration. In birds, the entire pelvis rocks on the axial skeleton to produce ventilation. Although the mechanisms of pelvic aspiration are very different in crocodilians and birds, it is unusual among vertebrates for the pelvic musculoskeletal system to play an active role in inspiration. This observation raises the possibility that the pelvic musculoskeletal system may have played an important role in the ventilation of basal archosaurs. Based on the mechanism of pelvic aspiration in crocodilians and the structure of gastralia in basal archosaurs, we suggest that an ischiotruncus muscle pulled the medial aspect of the gastralia caudally, and thereby helped to produce inspiration by increasing the volume of the cuirassal basket. The proposed mechanism of cuirassal breathing in non-avian theropods leads us to suggest that the phase relationship of the ventilatory and locomotor cycles in running theropods was the opposite of that observed in running birds. Furthermore, we suggest that the ventilatory cycle of flying pterosaurs was entrained to the locomotor cycle with a phase relationship that was the opposite of that observed during flight in modern birds.

Activity of reticulospinal neurons during locomotion in the freely behaving lamprey.

The reticulospinal (RS) system is the main descending system transmitting commands from the brain to the spinal cord in the lamprey. It is responsible for initiation of locomotion, steering, and equilibrium control. In the present study, we characterize the commands that are sent by the brain to the spinal cord in intact animals via the reticulospinal pathways during locomotion. We have developed a method for recording the activity of larger RS axons in the spinal cord in freely behaving lampreys by means of chronically implanted macroelectrodes. In this paper, the mass activity in the right and left RS pathways is described and the correlations of this activity with different aspects of locomotion are discussed. In quiescent animals, the RS neurons had a low level of activity. A mild activation of RS neurons occurred in response to different sensory stimuli. Unilateral eye illumination evoked activation of the ipsilateral RS neurons. Unilateral illumination of the tail dermal photoreceptors evoked bilateral activation of RS neurons. Water vibration also evoked bilateral activation of RS neurons. Roll tilt evoked activation of the contralateral RS neurons. With longer or more intense sensory stimulation of any modality and laterality, a sharp, massive bilateral activation of the RS system occurred, and the animal started to swim. This high activity of RS neurons and swimming could last for many seconds after termination of the stimulus. There was a positive correlation between the level of activity of RS system and the intensity of locomotion. An asymmetry in the mass activity on the left and right sides occurred during lateral turns with a 30% prevalence (on average) for the ipsilateral side. Rhythmic modulation of the activity in RS pathways, related to the locomotor cycle, often was observed, with its peak coinciding with the electromyographic (EMG) burst in the ipsilateral rostral myotomes. The pattern of vestibular response of RS neurons observed in the quiescent state, that is, activation with contralateral roll tilt, was preserved during locomotion. In addition, an inhibition of their activity with ipsilateral tilt was clearly seen. In the cases when the activity of individual neurons could be traced during swimming, it was found that rhythmic modulation of their firing rate was superimposed on their tonic firing or on their vestibular responses. In conclusion, different aspects of locomotor activity-initiation and termination, vigor of locomotion, steering and equilibrium control-are well reflected in the mass activity of the larger RS neurons.

Coding of locomotor phase in populations of neurons in rostral and caudal segments of the neonatal rat lumbar spinal cord.

Several experiments have demonstrated that rostral segments of the vertebrate lumbar spinal cord produce a rhythmic motor output more readily and of better quality than caudal segments. Here we examine how this rostrocaudal gradient of rhythmogenic capability is reflected in the spike activity of neurons in the rostral (L(2)) and caudal (L(5)) lumbar spinal cord of the neonatal rat. The spike activity of interneurons in the ventromedial cord, a region necessary for the production of locomotion, was recorded intracellularly with patch electrodes and extracellularly with tetrodes during pharmacologically induced locomotion. Both L(2) and L(5) neurons tended to be active in phase with their homologous ventral root. L(5) neurons, however, had a wider distribution of their preferred phases of activity throughout the locomotor cycle than L(2) neurons. The strength of modulation of the activity of individual L(2) neurons was also larger than that of L(5) neurons. These differences resulted in a stronger rhythmic signal from the L(2) neuronal population than from the L(5) population. These results demonstrate that the rhythmogenic capability of each spinal segment was reflected in the activity of interneurons located in the same segment. In addition to paralleling the rostrocaudal gradient of rhythmogenic capability, these results further suggest a colocalization of motoneurons and their associated interneurons involved in the production of locomotion.

Antidromic discharges in dorsal roots of decerebrate cats: I. Studies at rest and during fictive locomotion

Spontaneous rhythmic antidromic discharges have previously been recorded in proximal stumps of cut dorsal roots during locomotion (real and fictive). The goals of the present study were to elucidate (1) whether both orthodromic and antidromic discharges occur in the same dorsal root filament and (2) whether orthodromic discharges have an influence upon antidromic discharges of units in the same filament. Unitary activity was recorded in 70 uncut dorsal root filaments (L6-S1) in 15 decerebrate cats using bipolar Ag/AgCl electrodes. Spikes with similar wave shapes were considered to represent the activity of single units. Spike-triggered averaging (STA), local anaesthesia and transection of filaments were used to determine the direction of propagation of spikes. Spikes with different initial electrical polarities were found in most of the filaments and shown to propagate in opposite directions at rest and during fictive locomotion. On average, there were 38%±S.D. 23% antidromically discharging units per filament and their mean conduction velocity was 55 m/s±S.D. 25 m/s. After blocking orthodromic activity of the whole filament by a transection or local anesthesia applied distally to the recording site, changes were seen in the antidromic discharges of some units suggesting that spontaneous orthodromic discharges normally seen in the filament may influence the antidromic discharges of some units. Moreover, out of 27 antidromic units recorded during fictive locomotion, 12 were rhythmically modulated with peak discharges occurring in various parts of the locomotor cycle. We conclude that, in uncut dorsal roots, there is a normal coexistence of spontaneous orthodromic and antidromic discharges revealed by STA and that there is an interaction between spontaneous orthodromic and antidromic discharges.

Postcranial osteology and functional morphology of the Litopterna of Salla, Bolivia (Late Oligocene)

ABSTRACT Postcranial remains of litopterns of Salla, Bolivia are described and discussed in terms of their functional morphology. These elements are not associated with dental remains, but can be confidently referred to the family Macraucheniidae or Proterotheriidae. The macraucheniids (cf. Coniopternium spp.) have transversely stable joints indicating their specialization for parasagittal movements at these articulations (e.g., the humeral-antebrachial, cruxastragalar, and astragalonavicular joints). A profound suprapatellar fossa (“patellar pit”) of the femur is interpreted as having provided a place in which the patella could have become lodged, thus locking the knee-joint in the extended position. Facets on the cuboid and astragalus and their articulation in life position indicate that cuboastragular contact was present. This contact was slight, however, and may have occurred only during the loaded part of the locomotor cycle. A partial pes of a proterotheriid represents the first unambiguous evidence of this group at Salla and are the first Deseadan postcranial elements confidently referred to the Proterotheriinae. These elements indicate that the animal was tridactyl, but functionally monodactyl to the same degree as all Santacrucian proterotheriid genera except the “hypermonodactyl” Thoatherium. These litopterns and those to which they were compared record a great diversity of tarsal arrangements, such that simple characterizations of litoptern tarsals as being serial and “inadaptive” are unwarranted.

Water skating in the larvae of dixella aestivalis (Diptera) and hydrobius fuscipes (Coleoptera)

The kinematics of locomotion was investigated in the aquatic larvae of Dixella aestivalis and Hydrobius fuscipes with the aid of high-speed video recordings. Both insects are able to skate on the surface of the water using the dorso-apical tracheal gill as an adhesive organ or 'foot'. Progress relies on the variable adhesion of the foot between 'slide' and 'hold' periods of the locomotory cycle. The flexural body movements underlying skating in D. aestivalis can be derived directly from the figure-of-eight swimming mechanism used in underwater swimming. The latter is shown to be similar to figure-of-eight swimming in chironomid larvae. This study shows how the deployment of a 'foot' enables simple side-to-side flexural movements of the body to be converted into effective locomotion at the air-water interface.

Parallel control mechanisms underlying locomotor activity and sexual receptivity of the female German cockroach, Blattella germanica (L.)

We have studied the effects of ovaries, juvenile hormone (JH) and mating on locomotor activity and sexual receptivity of female German cockroaches. Our results indicate that locomotor activity and sexual receptivity are under the same control mechanisms. The ovary served as a negative masking factor for the locomotor circadian rhythm, but did not affect the frequency of locomotor activity. We conclude that JH controls the locomotor activity of females from the following evidence: (1) increasing locomotion of virgin females coincided with an increasing volume of the corpora allata; (2) allatectomy reduced female locomotion significantly; (3) after absorbing the JH analogue (fenoxycarb) through their tarsi, allatectomized females regained their high level of locomotor activity. Since the daily locomotor activity of allatectomized and ovari-allatectomized females changed cyclically with continuous (non-cyclic) contact of fenoxycarb, an unidentified factor which was independent of ovarian development is proposed to regulate cyclic locomotor activity. In addition to controlling the frequency of locomotor activity, JH was essential for the expression of the locomotor circadian rhythm because allatectomy abolished the circadian rhythm expressed in ovariectomized females. Mating significantly decreased the frequency of locomotor activity and the degree of sexual receptivity. The inhibitory effect of mating resulted from the transmission of a mating signal through the ventral nerve cord when sperm was transferred successfully. The mating experiments with allatectomized and ovariectomized females showed that JH was the major factor in regulating the expression of sexual receptivity.

Adaptive changes in responses to repeated locomotor perturbations in cerebellar patients.

This study examined the responses of cerebellar patients and a group of age- and sex-matched control subjects to repeated changes in treadmill speed in order to test whether cerebellar patients can adapt their gait to this type of perturbation and, if so, whether their responses are comparable to those of controls. While the subject walked on the treadmill, a perturbation consisting of a sudden slowing of the treadmill followed by a sudden increase back to the original speed was applied repeatedly at a specific time during the step cycle. Both the control subjects and cerebellar patients were able to compensate for the perturbations by minimizing their postural sway and changing step length. However, the nature of the compensatory changes in step length differed between these subject groups. Control subjects compensated for the perturbation by consistently using the same leg to initiate the response to the perturbation and by adapting a pattern of stepping such that the EMG characterizing the response occurred in a manner that was entrained to the timing of the normal locomotor cycle. In contrast, the patients, although undergoing modifications in step length, employed a much less consistent motor pattern from trial to trial than that of the normal subjects. An inconsistent pattern among their responses was apparent in both the analysis of stepping and in the EMG activity of the gastrocnemius and anterior tibial muscles. These results suggest that, although the cerebellar patients can adapt their behavior in response to locomotor perturbations, they do not establish a motor pattern comparable to that employed by normal subjects.

Effects of stimulating the reticular formation during fictive locomotion in lampreys

The effects of stimulating the reticular formation were studied during fictive locomotion in lampreys ( Ichthyomyzon unicuspis). The in vitro isolated preparation of the brainstem and spinal cord was used and fictive locomotion was induced by bath application of N-methyl- d-aspartate (NMDA; 50–100 μM). During different phases of the locomotor cycle, short trains of stimuli (10 pulses at 80–100 Hz; 10 μA) were delivered through glass-coated tungsten microelectrodes positioned within the middle rhombencephalic reticular nucleus (MRRN) and their effects were studied on ipsi- and contralateral ventral root locomotor discharges. Irrespective of the locomotor phase during which the stimulation train was delivered, a resetting effect occurred. It was characterized by a re-synchronization of the locomotor discharges with a constant latency for each ventral root on the ipsilateral side. The latency increased as the recorded root was located further caudally. This increase in latency was in the range of the phase lag observed between roots during control bouts of locomotion. These results suggest that reticulospinal neurones exert strong resetting effects on spinal locomotor networks. These effects may play a significant role with respect to changes of direction during swimming.

Trajectory of single primary afferents from the intercostal muscle spindle in the cat

Although the corticospinal tract is not essential for the production of the basic locomotor rhythm in cats, it does contribute to the regulation of locomotion, particularly in situations in which there is a requirement for precise control over paw placement or limb trajectory. Lesions of the dorsolateral funiculi at the low thoracic level (T13) that completely interrupted both the cortico- and rubrospinal pathways produced long-term deficits in locomotion on a level surface. These deficits included a paw-drag that was probably caused both by a loss of cortico- and rubrospinal input to motoneurones controlling distal muscles as well as by a change in the relative timing of muscles acting around the hip and knee. Smaller lesions produced similar deficits from which the cats recovered relatively quickly. Cats with the largest lesions of the dorsolateral funiculi were unable to modify their gait sufficiently to step over obstacles attached to the treadmill belt even 3–5 months postlesion. These results imply that the medial pathways, the reticulo- and vestibulospinal pathways, are unable to fully compensate for damage to the lateral pathways. Single unit recordings from identified pyramidal tract neurones (PTNs) within the hindlimb representation of the primary motor cortex (area 4) showed that a substantial proportion of neurones (67%) significantly increased their discharge frequency when the cats modified their gait to step over obstacles attached to the treadmill belt. Of those PTNs that showed increased activity during the swing phase, populations of neurones were activated at different times. A large proportion of PTNS discharged early in swing, in phase with knee flexors such as the semitendinosus. Others discharged slightly later, in phase with the activity of ankle flexors, such as tibialis anterior, while still others discharged at the end of swing, in phase with digit dorsiflexors, such as the extensor digitorum brevis. We suggest that different populations of cortical neurones may specifically modify the activity of selected groups of close synergistic muscles during different parts of the swing phase. We further suggest that these modifications are mediated, in part, by groups of interneurones that are involved in determining the base locomotor rhythm. This provides a means by which the changes specified by the descending signal from the motor cortex may be smoothly, and appropriately, incorporated into the locomotor cycle.