Gait acts as a gate for reflexes from the foot

2004 ◽  
Vol 82 (8-9) ◽  
pp. 715-722 ◽  
Author(s):  
J Duysens ◽  
C M Bastiaanse ◽  
B C.M Smits-Engelsman ◽  
V Dietz

During human gait, electrical stimulation of the foot elicits facilitatory P2 (medium latency) responses in TA (tibialis anterior) at the onset of the swing phase, while the same stimuli cause suppressive responses at the end of swing phase, along with facilitatory responses in antagonists. This phenomenon is called phase-dependent reflex reversal. The suppressive responses can be evoked from a variety of skin sites in the leg and from stimulation of some muscles such as rectus femoris (RF). This paper reviews the data on reflex reversal and adds new data on this topic, using a split-belt paradigm. So far, the reflex reversal in TA could only be studied for the onset and end phases of the step cycle, simply because suppression can only be demonstrated when there is background activity. Normally there are only 2 TA bursts in the step cycle, whereas TA is normally silent during most of the stance phase. To know what happens in the stance phase, one needs to have a means to evoke some background activity during the stance phase. For this purpose, new experiments were carried out in which subjects were asked to walk on a treadmill with a split-belt. When the subject was walking with unequal leg speeds, the walking pattern was adapted to a gait pattern resembling limping. The TA then remained active throughout most of the stance phase of the slow-moving leg, which was used as the primary support. This activity was a result of coactivation of agonistic and antagonistic leg muscles in the supporting leg, and represented one of the ways to stabilize the body. Electrical stimulation was given to a cutaneous nerve (sural) at the ankle at twice the perception threshold. Nine of the 12 subjects showed increased TA activity during stance phase while walking on split-belts, and 5 of them showed pronounced suppressions during the first part of stance when stimuli were given on the slow side. It was concluded that a TA suppressive pathway remains open throughout most of the stance phase in the majority of subjects. The suggestion was made that the TA suppression increases loading of the ankle plantar flexors during the loading phase of stance.Key words: human gait, cutaneous reflexes, sural nerve, tibialis anterior, split belt, reflex reversal.

1990 ◽  
Vol 63 (5) ◽  
pp. 1118-1127 ◽  
Author(s):  
L. Villanueva ◽  
K. D. Cliffer ◽  
L. S. Sorkin ◽  
D. Le Bars ◽  
W. D. Willis

1. Recordings were made in anesthetized monkeys from neurons in the medullary reticular formation (MRF) caudal to the obex. Responses of 19 MRF neurons to mechanical, thermal, and/or electrical stimulation were examined. MRF neurons exhibited convergence of nociceptive cutaneous inputs from widespread areas of the body and face. 2. MRF neurons exhibited low levels of background activity. Background activity increased after periods of intense cutaneous mechanical or thermal stimulation. Nearly all MRF neurons tested failed to respond to heterosensory stimuli (flashes, whistle sounds), and none responded to joint movements. 3. MRF neurons were excited by and encoded the intensity of noxious mechanical stimulation. Responses to stimuli on contralateral limbs were greater than those to stimuli on ipsilateral limbs. Responses were greater to stimuli on the forelimbs than to stimuli on the hindlimbs. 4. MRF neurons responded to noxious thermal stimulation (51 degrees C) of widespread areas of the body. Mean responses from stimulation at different locations were generally parallel to those for noxious mechanical stimulation. Responses increased with intensity of noxious thermal stimulation (45-50 degrees C). 5. MRF neurons responded with one or two peaks of activation to percutaneous electrical stimulation applied to the limbs, the face, or the tail. The differences in latency of responses to stimulating two locations along the tail suggested that activity was elicited by activation of peripheral fibers with a mean conduction velocity in the A delta range. Stimulation of the contralateral hindlimb elicited greater responses, with lower thresholds and shorter latencies, than did stimulation of the ipsilateral hindlimb. 6. Electrophysiological properties of monkey MRF neurons resembled those of neurons in the medullary subnucleus reticularis dorsalis (SRD) in the rat. Neurons in the caudal medullary reticular formation could play a role in processing nociceptive information. Convergence of nociceptive cutaneous input from widespread areas of the body suggests that MRF neurons may contribute to autonomic, affective, attentional, and/or sensory-motor processes related to pain.


1996 ◽  
Vol 75 (3) ◽  
pp. 1126-1137 ◽  
Author(s):  
G. W. Hiebert ◽  
P. J. Whelan ◽  
A. Prochazka ◽  
K. G. Pearson

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.


Author(s):  
J.F. Yang ◽  
J. Fung ◽  
M. Edamura ◽  
R. Blunt ◽  
R.B. Stein ◽  
...  

ABSTRACT:Hoffmann (H) reflexes were elicited from the soleus muscle during treadmill walking in 21 spastic paretic patients. The soleus and tibialis anterior muscles were reciprocally activated during walking in most patients, much like that observed in healthy individuals. The pattern of H-reflex modulation varied considerably between patients, from being relatively normal in some patients to a complete absence of modulation in others. The most common pattern observed was a lack of H-reflex modulation through the stance phase and slight depression of the reflex in the swing phase, considerably less modulation than that of normal subjects under comparable walking conditions. The high reflex amplitudes during periods of the step cycle such as early stance seems to be related to the stretch-induced large electromyogram bursts in the soleus in some subjects. The abnormally active reflexes appear to contribute to the clonus encountered during walking in these patients. In three patients who were able to walk for extended periods, the effect of stimulus intensity was examined. Two of these patients showed a greater degree of reflex modulation at lower stimulus intensities, suggesting that the lack of modulation observed at higher stimulus intensities is a result of saturation of the reflex loop. In six other patients, however, no reflex modulation could be demonstrated even at very low stimulus intensities.


1997 ◽  
Vol 01 (02) ◽  
pp. 95-109 ◽  
Author(s):  
W. Herzog ◽  
T. R. Leonard

The dynamic properties of the cat soleus muscle were studied in freely walking animal preparations. The force and EMG responses of the soleus following supramaximal, ins tants of the step cycle. The sensorimotor interactions of soleus with the medial head of the gastrocnemius (a functional agonist of the soleus at the ankle) and the tibialis anterior (a functional antagonist of soleus at the ankle) were studied by measuring their force and EMG responses following the artifical stimulation of the soleus nerve. Supramaximal nerve stimulation showed distinct increases in the soleus forces during the entire swing phase and the second part (after peak forces had been reached) of the stance phase. Soleus forces could only be increased slightly in the first part of stance (from paw contact to peak force). These results suggest that force production of the soleus is virtually maximal during the early phases of stance but is submaximal for the remainder of the step cycle. Forces and EMGs of the medial gastrocnemius muscle were affected by the soleus nerve stimulation only in the latter part of the swing phase. In these cases, the force and EMG of the medial gastrocnemius were reduced significantly for the step cycle following the perturbation. The active force production of soleus during late swing causes an inhibition of medial gastrocnemius activity and force. Forces and EMGs of the tibialis anterior muscle were always affected by the soleus nerve stimulation during the swing phase of the step cycle. In these case, the force EMG of the medial gastrocnemius were reduced significantly for the step cycle following the perturbation. The active force production of soleus during late swing causes an inhibition of medial gastrocnemius activity and force. Forces and EMGs of the tibialis anterior muscle were always affected by the soleus nerve stimulation during the swing phase of the step cycle. In these instances, forces and EMGs of the tibialis anterior were significantly increased compared to step cycles preceding or following the perturbation. Part of the force enhancement is caused by the stretch of the activated tibialis anterior by the soleus, and part of the enhancement is caused by reflex activation. No effects on forces or EMGs of the tibialis anterior were observed when the soleus nerve stimulation showed its effects during the stance phase of the step cycle. The results of theis study suggest that the magnitude and the quality of ensorimotor interactions of soleus with medial gastrocnemius and tibialis anterior depend on the phase of the step cycle. The strongest interactions appear to exist during the swing phase; no observable interactions were found during stance.


2003 ◽  
Vol 90 (5) ◽  
pp. 2850-2861 ◽  
Author(s):  
Carlos Haridas ◽  
E. Paul Zehr

It has been shown that stimulation of cutaneous nerves innervating the hand (superficial radial, SR) and foot (superficial peroneal, SP) elicit widespread reflex responses in many muscles across the body. These interlimb reflex responses were suggested to be functionally relevant to assist in motor coordination between the arms and legs during motor tasks such as walking. The experiments described in this paper were conducted to test the hypothesis that interlimb reflexes were phase-dependently modulated and produced functional kinematic changes during locomotion. Subjects walked on a treadmill while electromyographic (EMG) activity was collected continuously from all four limbs, and kinematic recordings were made of angular changes across the ankle, knee, elbow, and shoulder joints. Cutaneous reflexes were evoked by delivering trains of electrical stimulation pseudorandomly to the SP nerve or SR nerves in separate trials. Reflexes were phase-averaged according to the time of occurrence in the step cycle, and phasic amplitudes and latencies were calculated. For both nerves, significant phase-dependent modulation (including reflex reversals) of interlimb cutaneous reflex responses was seen in most muscles studied. Both SR and SP nerve stimulation resulted in significant alteration in ankle joint kinematics. The results suggest coordinated and functionally relevant reflex pathways from the SP and SR nerves onto motoneurons innervating muscles in nonstimulated limbs during walking, thus extending observations from the cat to that of the bipedal human.


2016 ◽  
Vol 116 (1) ◽  
pp. 183-190 ◽  
Author(s):  
Tsuyoshi Nakajima ◽  
Shinya Suzuki ◽  
Genki Futatsubashi ◽  
Hiroyuki Ohtsuska ◽  
Rinaldo A. Mezzarane ◽  
...  

During walking, cutaneous reflexes in ankle flexor muscle [tibialis anterior (TA)] evoked by tibial nerve (TIB) stimulation are predominantly facilitatory at early swing phase but reverse to suppression at late swing phase. Although the TIB innervates a large portion of the skin of the foot sole, the extent to which specific foot-sole regions contribute to the reflex reversals during walking remains unclear. Therefore, we investigated regional cutaneous contributions from discrete portions of the foot sole on reflex reversal in TA following TIB stimulation during walking. Summation effects on reflex amplitudes, when applying combined stimulation from foot-sole regions with TIB, were examined. Middle latency responses (MLRs; 70–120 ms) after TIB stimulation were strongly facilitated during the late stance to mid-swing phases and reversed to suppression just before heel (HL) strike. Both forefoot-medial (f-M) and forefoot-lateral stimulation in the foot sole induced facilitation during stance-to-swing transition phases, but HL stimulation evoked suppression during the late stance to the end of swing phases. At the stance-to-swing transition, a summation of MLR amplitude occurred only for combined f-M&TIB stimulation. However, the same was not true for the combined HL&TIB stimulation. At the swing-to-stance transition, there was a suppressive reflex summation only for HL&TIB stimulation. In contrast, this summation was not observed for the f-M&TIB stimulation. Our results suggest that reflex reversals evoked by TIB stimulation arise from distinct reflex pathways to TA produced by separate afferent populations innervating specific regions of the foot sole.


1995 ◽  
Vol 198 (7) ◽  
pp. 1589-1601 ◽  
Author(s):  
F Kuenzi ◽  
M Burrows

The hair plate proprioceptors at the thoraco-coxal joint of insect limbs provide information about the movements of the most basal joint of the legs. The ventral coxal hair plate of a middle leg consists of group of 10-15 long hairs (70 microns) and 20-30 short hairs (30 microns). The long hairs are deflected by the trochantin as the leg is swung forward during the swing phase of walking, and their sensory neurones respond phasically during an imposed deflection and tonically if the deflection is maintained. Selective stimulation of the long hairs elicits a resistance reflex that rotates the coxa posteriorly and is similar to that occurring at the transition from the swing to the stance phase of walking. The motor neurones innervating the posterior rotator and adductor coxae muscles are excited, and those to the antagonistic anterior rotator muscle are inhibited. By contrast, selective stimulation of the short hairs leads only to a weak inhibition of the anterior rotator. The excitatory effects of the long hairs are mediated, in part, by direct connections between their sensory neurones and particular motor neurones. A spike in a sensory neurone elicits a short-latency depolarising postsynaptic potential (PSP) in posterior rotator and adductor motor neurones whose amplitude is enhanced by hyperpolarising current injected into the motor neurone. When the calcium in the saline is replaced with magnesium, the amplitude of the PSP is reduced gradually, and not abruptly as would be expected if an interneurone were interposed in the pathway. Several sensory neurones from long hairs converge to excite an individual motor neurone, evoking spikes in some motor neurones. The projections of the sensory neurones overlap with some of the branches of the motor neurones in the lateral association centre of the neuropile. It is suggested that these pathways would limit the extent of the swing phase of walking and contribute to the switch to the stance phase in a negative feedback loop that relieves the excitation of the hairs by rotating the coxa backwards.


1990 ◽  
Vol 148 (1) ◽  
pp. 113-128 ◽  
Author(s):  
U. W. E. MÜLLER ◽  
FRANÇOIS CLARAC

1. The effects of cyclic electrical stimulation of the dactyl sensory nerve (DN) on the walking pattern of rock lobsters were examined at the two crucial points within the step cycle: the anterior extreme position (AEP) and the posterior extreme position (PEP). 2. Stimulation during the occurrence of the PEP affected neither the movement pattern of the stimulated leg itself nor that of the ipsilateral adjacent legs. 3. Stimulation of the same intensity during the occurrence of the AEP interrupted the oscillation of the stimulated leg and affected the phase relationships of the ipsilateral adjacent legs. 4. The possibility that indirect influences were mediated by coupling to the substratum can be excluded. Neuronal connections may therefore exist between the funnel canal organs (FCO) of a single leg and the motor output of the adjacent legs. The discussion deals with whether the described channels alone are able to fulfil the requirements of a ‘coordinating mechanism’ as described in the literature.


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