Contribution of the Motor Cortex to the Structure and the Timing of Hindlimb Locomotion in the Cat: A Microstimulation Study

2005 ◽  
Vol 94 (1) ◽  
pp. 657-672 ◽  
Author(s):  
Frédéric Bretzner ◽  
Trevor Drew
Keyword(s):  
Motor Cortex ◽  
Muscle Activity ◽  
Pyramidal Tract ◽  
Cycle Duration ◽  
Step Cycle ◽  
Standard Glass ◽  
Level Of Activity ◽  
Flexor Muscles ◽  
Multijoint Movements ◽  
Stimulation Of

We used microstimulation to examine the contribution of the motor cortex to the structure and timing of the hindlimb step cycle during locomotion in the intact cat. Stimulation was applied to the hindlimb representation of the motor cortex in 34 sites in three cats using either standard glass-insulated microelectrodes (16 sites in 1 cat) or chronically implanted microwire electrodes (18 sites in 2 cats). Stimulation at just suprathreshold intensities with the cat at rest produced multijoint movements at a majority of sites (21/34, 62%) but evoked responses restricted to a single joint, normally the ankle, at the other 13/34 (38%) sites. Stimulation during locomotion generally evoked larger responses than the same stimulation at rest and frequently activated additional muscles. Stimulation at all 34 sites evoked phase-dependent responses in which stimulation in swing produced transient increases in activity in flexor muscles while stimulation during stance produced transient decreases in activity in extensors. Stimulation with long (200 ms) trains of stimuli in swing produced an increased level of activity and duration of flexor muscles without producing changes in cycle duration. In contrast, stimulation during stance decreased the duration of the extensor muscle activity and initiated a new and premature period of swing, resetting the step cycle. Stimulation of the pyramidal tract in two of these three cats as well as in two additional ones produced similar effects. The results show that the motor cortex is capable of influencing hindlimb activity during locomotion in a similar manner to that seen for the forelimb.

Effects of Red Nucleus Microstimulation on the Locomotor Pattern and Timing in the Intact Cat: A Comparison With the Motor Cortex

1999 ◽  
Vol 81 (5) ◽  
pp. 2297-2315 ◽  
Author(s):  
Marie-Josée Rho ◽  
Sylvain Lavoie ◽  
Trevor Drew
Keyword(s):  
Motor Cortex ◽  
Red Nucleus ◽  
Emg Activity ◽  
Cycle Duration ◽  
Step Cycle ◽  
Rubrospinal Tract ◽  
Locomotor Pattern ◽  
Train Duration ◽  
Flexor Muscles ◽  
Cathodal Current

Effects of red nucleus microstimulation on the locomotor pattern and timing in the intact cat: a comparison with the motor cortex. To determine the extent to which the rubrospinal tract is capable of modifying locomotion in the intact cat, we applied microstimulation (cathodal current, 330 Hz; pulse duration 0.2 ms; maximal current, 25 μA) to the red nucleus during locomotion. The stimuli were applied either as short trains (33 ms) of impulses to determine the capacity of the rubrospinal tract to modify the level of electromyographic (EMG) activity in different flexors and extensors at different phases of the step cycle or as long trains (200 ms) of pulses to determine the effect of the red nucleus on cycle timing. Stimuli were also applied with the cat at rest (33-ms train). This latter stimulation evoked short-latency (average = 11.8–19.0 ms) facilitatory responses in all of the physiological flexor muscles of the forelimb that were recorded; facilitatory responses were also common in the elbow extensor, lateral head of triceps but were rare in the physiological wrist and digit extensor, palmaris longus. Responses were still evoked in most muscles when the current was decreased to near threshold (3–10 μA). Stimulation during locomotion with the short trains of stimuli evoked shorter-latency (average = 6.0–12.5 ms) facilitatory responses in flexor muscles during the swing phase of locomotion and, except in the case of the extensor digitorum communis, evoked substantially smaller responses in stance. The same stimuli also evoked facilitatory responses in the extensor muscles during swing and produced more complex effects involving both facilitation and suppression in stance. Increasing the duration of the train to 200 ms modified the amplitude and duration of the EMG activity of both flexors and extensors but had little significant effect on the cycle duration. In contrast, whereas stimulation of the motor cortex with short trains of stimuli during locomotion had very similar effects to that of the red nucleus, increasing the train duration to 200 ms frequently produced a marked reset of the step cycle by curtailing stance and initiating a new period of swing. The results suggest that whereas both the motor cortex and the red nucleus have access to the interneuronal circuits responsible for controlling the structure of the EMG activity in the step cycle, only the motor cortex has access to the circuits responsible for controlling cycle timing.

scholarly journals Inhibitory influence of the ipsilateral motor cortex on responses to stimulation of the human cortex and pyramidal tract

1998 ◽  
Vol 510 (1) ◽  
pp. 249-259 ◽  
Author(s):  
Christian Gerloff ◽  
Leonardo G. Cohen ◽  
Mary Kay Floeter ◽  
Robert Chen ◽  
Brian Corwell ◽  
...  
Keyword(s):  
Motor Cortex ◽  
Pyramidal Tract ◽  
Inhibitory Influence ◽  
Human Cortex ◽  
Stimulation Of

Contribution of hind limb flexor muscle afferents to the timing of phase transitions in the cat step cycle

1996 ◽  
Vol 75 (3) ◽  
pp. 1126-1137 ◽  
Author(s):  
G. W. Hiebert ◽  
P. J. Whelan ◽  
A. Prochazka ◽  
K. G. Pearson
Keyword(s):  
Electrical Stimulation ◽  
Stance Phase ◽  
Flexor Muscle ◽  
Step Cycle ◽  
Locomotor Rhythm ◽  
Ia Afferents ◽  
Group I ◽  
Flexor Muscles ◽  
Group Ii ◽  
Stimulation Of

1. In this investigation, we tested the hypothesis that muscle spindle afferents signaling the length of hind-leg flexor muscles are involved in terminating extensor activity and initiating flexion during walking. The hip flexor muscle iliopsoas (IP) and the ankle flexors tibialis anterior (TA) and extensor digitorum longus (EDL) were stretched or vibrated at various phases of the step cycle in spontaneously walking decerebrate cats. Changes in electromyogram amplitude, duration, and timing were then examined. The effects of electrically stimulating group I and II afferents in the nerves to TA and EDL also were examined. 2. Stretch of the individual flexor muscles (IP, TA, or EDL) during the stance phase reduced the duration of extensor activity and promoted the onset of flexor burst activity. The contralateral step cycle also was affected by the stretch, the duration of flexor activity being shortened and extensor activity occurring earlier. Therefore, stretch of the flexor muscles during the stance phase reset the locomotor rhythm to flexion ipsilaterally and extension contralaterally. 3. Results of electrically stimulating the afferents from the TA and EDL muscles suggested that different groups of afferents were responsible for the resetting of the step cycle. Stimulation of the TA nerve reset the locomotor step cycle when the stimulus intensity was in the group II range (2-5 xT). By contrast, stimulation of the EDL nerve generated strong resetting of the step cycle in the range of 1.2-1.4 xT, where primarily the group Ia afferents from the muscle spindles would be activated. 4. Vibration of IP or EDL during stance reduced the duration of the extensor activity by similar amounts to that produced by muscle stretch or by electrical stimulation of EDL at group Ia strengths. This suggests that the group Ia afferents from IP and EDL are capable of resetting the locomotor pattern generator. Vibration of TA did not affect the locomotor rhythm. 5. Stretch of IP or electrical stimulation of TA afferents (5 xT) during the flexion phase did not change the duration of the flexor activity. Stimulation of the EDL nerve at 1.8-5 xT during flexion increased the duration of the flexor activity. In none of our preparations did we observe resetting to extension when the flexor afferents were activated during flexion. 6. We conclude that as the flexor muscles lengthen during the stance phase of gait, their spindle afferents (group Ia afferents for EDL and IP, group II afferents for TA) act to inhibit the spinal center generating extensor activity thus facilitating the initiation of swing.

scholarly journals Studies on the Corticospinal Control of Human Walking. I. Responses to Focal Transcranial Magnetic Stimulation of the Motor Cortex

1999 ◽  
Vol 81 (1) ◽  
pp. 129-139 ◽  
Author(s):  
Charles Capaday ◽  
Brigitte A. Lavoie ◽  
Hugues Barbeau ◽  
Cyril Schneider ◽  
Mireille Bonnard
Keyword(s):  
Motor Cortex ◽  
Magnetic Stimulation ◽  
Stance Phase ◽  
Corticospinal Tract ◽  
Human Walking ◽  
Step Cycle ◽  
Motor Circuits ◽  
Significant Difference ◽  
Stimulation Of

Capaday, Charles, Brigitte A. Lavoie, Hugues Barbeau, Cyril Schneider, and Mireille Bonnard. Studies on the corticospinal control of human walking. I. Responses to focal transcranial magnetic stimulation of the motor cortex. J. Neurophysiol. 81: 129–139, 1999. Experiments were done to determine the extent to which the corticospinal tract is linked with the segmental motor circuits controlling ankle flexors and extensors during human walking compared with voluntary motor tasks requiring attention to the level of motor activity. The motor cortex was activated transcranially using a focal magnetic stimulation coil. For each subject, the entire input-output (I-O) curve [i.e., the integral of the motor evoked-potential (MEP) versus stimulus strength] was measured during a prescribed tonic voluntary contraction of either the tibialis anterior (TA) or the soleus. Similarly, I-O curves were measured in the early part of the swing phase, or in the early part of the stance phase of walking. The I-O data points were fitted by the Boltzmann sigmoidal function, which accounted for ≥80% of total data variance. There was no statistically significant difference between the I-O curves of the TA measured during voluntary ankle dorsiflexion or during the swing phase of walking, at matched levels of background electromyographic (EMG) activity. Additionally, there was no significant difference in the relation between the coefficient of variation and the amplitude of the MEPs measured in each task, respectively. In comparison, during the stance phase of walking the soleus MEPs were reduced on average by 26% compared with their size during voluntary ankle plantarflexion. Furthermore, during stance the MEPs in the inactive TA were enhanced relative to their size during voluntary ankle plantarflexion and in four of six subjects the TA MEPs were larger than those of the soleus. Finally, stimulation of the motor cortex at various phases of the step cycle did not reset the cycle. The time of the next step occurred at the expected moment, as determined from the phase-resetting curve. One interpretation of this result is that the motor cortex may not be part of the central neural system involved in timing the motor bursts during the step cycle. We suggest that during walking the corticospinal tract is more closely linked with the segmental motor circuits controlling the flexor, TA, than it is with those controlling the extensor, soleus. However, during voluntary tasks requiring attention to the level of motor activity, it is equally linked with the segmental motor circuits of ankle flexors or extensors.

Activity of the Motor Cortex During Scratching

2006 ◽  
Vol 95 (2) ◽  
pp. 753-765 ◽  
Author(s):  
Mikhail G. Sirota ◽  
Galina A. Pavlova ◽  
Irina N. Beloozerova
Keyword(s):  
Motor Cortex ◽  
Hind Limb ◽  
Pyramidal Tract ◽  
Average Depth ◽  
Simple Correlation ◽  
Population Activity ◽  
Ankle Muscles ◽  
Pyramidal Tract Neurons ◽  
Awake Cats ◽  
Stimulation Of

In awake cats sitting with the head restrained, scratching was evoked using stimulation of the ear. Cats scratched the shoulder area, consistently failing to reach the ear. Kinematics of the hind limb movements and the activity of ankle muscles, however, were similar to those reported earlier in unrestrained cats. The activity of single neurons in the hind limb representation of the motor cortex, including pyramidal tract neurons (PTNs), was examined. During the protraction stage of the scratch response, the activity in 35% of the neurons increased and in 50% decreased compared with rest. During the rhythmic stage, the motor cortex population activity was approximately two times higher compared with rest, because the activity of 53% of neurons increased and that of 33% decreased in this stage. The activity of 61% of neurons was modulated in the scratching rhythm. The average depth of frequency modulation was 12.1 ± 5.3%, similar to that reported earlier for locomotion. The phases of activity of different neurons were approximately evenly distributed over the scratch cycle. There was no simple correlation between resting receptive field properties and the activity of neurons during the scratch response. We conclude that the motor cortex participates in both the protraction and the rhythmic stages of the scratch response.

Functional organization within the medullary reticular formation of the intact unanesthetized cat. III. Microstimulation during locomotion

1991 ◽  
Vol 66 (3) ◽  
pp. 919-938 ◽  
Author(s):  
T. Drew
Keyword(s):  
Reticular Formation ◽  
Muscle Activity ◽  
Functional Organization ◽  
Vastus Lateralis ◽  
Medial Longitudinal Fasciculus ◽  
Step Cycle ◽  
Medullary Reticular Formation ◽  
Extensor Muscles ◽  
Flexor Muscles ◽  
Excitatory Responses

1. This article presents the results from stimulation in 21 loci within the medullary reticular formation (MRF; between 0.5 and 2.5 mm from the midline) and in 5 loci in the medial longitudinal fasciculus (MLF) of four intact, unanesthetized cats during locomotion. Stimulus trains (11 pulses, 0.2-ms duration, 330 Hz, stimulus strength 35 microA) were applied at those loci in each track at which the most widespread effects in each of the four limbs were obtained with the cat at rest. Electromyograms were recorded from flexor and extensor muscles of each limb. 2. As previously reported, stimulation with the cat at rest generally evoked brief, short-latency, twitch responses in both flexor and extensor muscles of more than one limb. In contrast, stimulation during locomotion evoked a more complex pattern of activity in which responses were normally evoked in one or other of the muscle pairs and incorporated into the locomotor pattern. 3. In the majority of sites, the stimulation evoked excitatory responses in the flexor muscles of each of the four limbs during that period of the step cycle in which each respective muscle was naturally active; stimulation in the stance phase of locomotion, although less effective, was also capable of producing responses in these muscles. All three ipsilateral extensor muscles studied [long and lateral heads of triceps and vastus lateralis (Tri, TriL, and VL, respectively)] were normally inhibited during their phase of muscle activity, although excitatory responses were occasionally seen. Responses in the contralateral (co) Tri were invariably excitatory and were largest during the period of muscle activity, whereas responses during the period of activity of the coVL were mixed, with both excitatory and inhibitory responses being seen from any one locus. 4. Excitatory responses were normally largest when stimulation was applied during the time that the muscle was active during the locomotor cycle. Responses evoked at times when the muscle was inactive were sometimes larger than those evoked with the animal at rest; such responses were most commonly seen in the hindlimb flexors and in the coVL. 5. In both flexors and extensors of each of the four limbs, the latency of the responses was greatest when the cat was at rest and least for stimuli given during the period of activity of the respective muscle. Average latencies during the period of muscle activity ranged from a minimum of 9.0 +/- 2.6 (SD) ms for inhibitory responses in the ipsilateral Tri and TriL to a maximum of 17.1 +/- 3.0 ms for the responses evoked in the ipsilateral semitendinosus.(ABSTRACT TRUNCATED AT 400 WORDS)

Control of Locomotor Cycle Durations

2005 ◽  
Vol 94 (2) ◽  
pp. 1057-1065 ◽  
Author(s):  
S. Yakovenko ◽  
D. A. McCrea ◽  
K. Stecina ◽  
A. Prochazka
Keyword(s):  
Sensory Input ◽  
Cycle Duration ◽  
Fictive Locomotion ◽  
Experimental Phase ◽  
Step Cycle ◽  
Phase Duration ◽  
Two Parameters ◽  
Lower Background ◽  
Sensitivity Gain ◽  
Stimulation Of

In intact animals and humans, increases in locomotor speed are usually associated with decreases in step cycle duration. Most data indicate that the locomotor central pattern generator (CPG) shortens cycle duration mainly by shortening the durations of extensor rather than flexor phases of the step cycle. Here we report that in fictive locomotion elicited by electrical stimulation of the midbrain locomotor region (MLR) in the cat, spontaneous variations in cycle duration were due more to changes in flexor rather than extensor phase durations in 22 of 31 experiments. The locomotor CPG is therefore not inherently extensor- or flexor-biased. We coined the term “dominant” to designate the phase (flexion or extension) showing the larger variation. In a simple half-center oscillator model, experimental phase duration plots were fitted well by adjusting two parameters that corresponded to background drive (“bias”) and sensitivity (“gain”) of the oscillator's timing elements. By analogy we argue that variations in background drive to the neural timing elements of the CPG could produce larger variations in phase duration in the half-center receiving the lower background drive, i.e., background drive may determine which half-center is dominant. The fact that data from normal cats were also fitted well by the model indicates that sensory input and central drive combine to determine locomotor phase durations. We conclude that there is a considerable flexibility in the control of phase durations in MLR-induced fictive locomotion. We posit that this may be explained by changes in background excitation of neural timing elements in the locomotor CPG.

scholarly journals Focal Increases of White Matter Glucose Utilization Produced by Electrical Stimulation of Rat Motor Cortex

1983 ◽  
Vol 3 (1) ◽  
pp. 67-70 ◽  
Author(s):  
Frank R. Sharp ◽  
Sherzad Bzorgchami ◽  
Thomas Kilduff
Keyword(s):  
Electrical Stimulation ◽  
White Matter ◽  
Motor Cortex ◽  
Glucose Utilization ◽  
Pyramidal Tract ◽  
Internal Capsule ◽  
Local Cerebral Glucose Utilization ◽  
The Right ◽  
Deoxyglucose Method ◽  
Stimulation Of

The right motor cortex was electrically stimulated in adult, awake rats for 45 min. Local cerebral glucose utilization (LCGU) was measured in white matter pathways with the (14C)-2-deoxyglucose method. Stimulation increased LCGU in focal regions of the right internal capsule to 51.3 μmol/100 g/min, compared to 39.8 on the control left side. Stimulation also increased LCGU in the right, medial pontine pyramidal tract to 36.2 μmol/100 g/min, compared with 27.3 on the control left side. The data demonstrate that electrical stimulation of motor cortex neurons increases LCGU 30 to 40% in the efferent myelinated axons of those neurons.

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