Phase-dependent responses in locomotor muscles of walking man

Bilateral reflex responses in locomotor muscles were studied in normal human subjects walking on a treadmill. Reflex responses were elicited in response to a momentary resistance applied to one leg. The responses were recorded electromyographically from superficial muscles of both lower limbs: the vastus lateralis, gluteus medius and tibialis anterior. A four-channel storage oscilloscope displayed a quartet record which consisted of phases of the walking cycle and patterns of EMG activity. Resistance and response data were collected for comparison. The appearance of reflex responses was found to be conditioned. The following two observations provide the phase-dependent characteristics of those responses: muscles of the ipsilateral limb elicited responses throughout the walking cycle regardless of whether the muscles were active or silent and muscles of the contralateral limb produced responses that depended on the point at which resistance occurred during the walking cycle. Discusion follows concerning inhibition of the response that may be responsible for this phase-dependent phenomenon.     

Different fusimotor reflexes from the ipsi- and contralateral hind limbs of the cat assessed in the same primary muscle spindle afferents

Experiments were performed in forty-five cats anaesthetized with alpha-chloralose. The aim of the study was to investigate a sample of primary muscle spindle afferents from triceps muscle with respect to their fusimotor reflex control from ipsi- as well as contralateral hind limb. Primary muscle spindle afferents of the triceps surae muscle were recorded from the mean rate of firing and the modulation of the afferent response to sinusoidal stretching of the triceps surae muscle was determined. Test measurements were made during tonic stretch of the ipsilateral PBSt, contralateral PBSt, contralateral triceps muscle or during extension of the intact contralateral hind limb. Control measurements were made with ipsi- and contralateral PBSt as well as contralateral triceps muscles relaxed and with contralateral hind limb in resting position. The occurrence and types of fusimotor effects were assessed by comparing test to control responses. The main finding of the present investigation was the great variability in type and size of the fusimotor effects evoked by different ipsi- and contralateral reflex stimuli. Both ipsi- and contralateral stimulations gave rise to predominantly dynamic, predominantly static or mixed static and dynamic fusimotor reflexes. In the same preparation, a given reflex stimulus often caused different reflex responses in different triceps surae primary spindle afferents. In the same afferent unit, different reflex stimuli usually produced fusimotor effects which differed from each other in type and/or size. In general, contralateral whole limb extension and stretch of contralateral PBSt muscles were more potent as reflex stimuli than stretch of the ipsilateral PBSt muscle. Stretch of the contralateral triceps surae muscle was, but for a few afferent units, ineffective as reflexogenic stimulus. It is concluded that the individualized receptive profiles of the primary muscle spindle afferents, which have been postulated in earlier investigations where the effects of different stimuli have been investigated on different cell populations, still seems to hold good when the stimuli are tested on the same units. The individuality of the receptive profiles of gamma-motoneurones is discussed in relation to different motor control hypotheses.     

The influence of vibration on spinal alpha-motoneurons excitability in static conditions and during evoked stepping in human

In healthy human the excitability of spinal alpha-motoneurons under application of vibrostimulation (20-60 Hz) to different leg muscles was investigated both in stationary condition and during stepping movements caused by vibration in the condition of suspended leg. In 15 subjects the amplitude of H-reflex were compared under vibration of rectus femoris (RF) and biceps femoris (BF) muscles of left leg as well during vibration of rectus femoris of contralateral, motionless leg in three spatial positions: upright, supine and on right side of body with suspended left leg. In dynamic conditions the amount of H-reflex was compared during evoked and voluntary stepping at 8 intervals of step cycle. In all body positions the vibration of each ipsilateral leg muscles caused significant suppression of H-reflex, this suppression was more prominent in the air-stepping conditions. The vibration of contralateral leg RF muscle had a weak influence on the amplitude of H-reflex. In 7 subjects the muscle vibration of ipsilateral and contralateral legs generated stepping movements. During evoked "air-stepping" H-reflex had different amplitudes in different phases of step cycle. At the same time the differences between responses under voluntary and non-voluntary stepping were revealed only in stance phase. Thus, different degree of H-reflex suppression by vibration under different body position in space depends on, it seems to be, from summary afferent inflows to spinal cord interneurons, which participate in regulation of posture and locomotion. Seemingly, the increasing of spinal cord neurons excitability occurs under involuntary air-stepping in swing phase, which is necessary for activation of locomotor automatism under unloading leg conditions.     

Influence of vibration on the excitability of spinal α-motoneurons under static conditions and during evoked stepping in humans

The excitability of spinal α-motoneurons in healthy humans was investigated with vibrostimulation (20–60 Hz) applied to different groups of muscles both under stationary conditions and during vibration-evoked stepping movements with leg suspension. In 15 subjects, the H-reflex amplitude was compared under the conditions of vibration of the left leg quadriceps femoris (QFM) or biceps femoris (BFM) muscle, as well as under the conditions of vibration of the contralateral, motionless leg QFM muscle in three spatial positions of the body: upright, supine, and lying on the side with the left leg suspended. Under dynamic conditions, the H-reflex value was compared during evoked and voluntary steppings at eight intervals of the step cycle. In all body positions, the vibration of each ipsilateral leg muscle caused a significant H-reflex suppression, this suppression being more prominent under the air-stepping conditions. The vibration of the contralateral leg QFM had weak influence on the H-reflex amplitude. In seven subjects, the vibration of the ipsilateral and contralateral leg muscles generated stepping movements. During vibration-evoked air-stepping, the H-reflex had different amplitudes in different phases of the step cycle. At the same time, the differences between responses under voluntary and involuntary stepping conditions were revealed only in the step cycle phase corresponding to the stance phase. Thus, the different degrees of the H-reflex suppression by vibration in different spatial positions of the body seem to depend on the summary afferent inflows to the spinal cord interneurons involved in the regulation of locomotion and posture. Apparently, an increase in the spinal cord neuronal excitability, which is necessary for activating locomotor automatism under the leg unloading conditions, occurs during evoked air-stepping in the swing phase.     

Gait acts as a gate for reflexes from the foot

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.     

On the reflex coactivation of ankle flexor and extensor muscles induced by a sudden drop of support surface during walking in humans.

Recent studies have revealed that the stretch reflex responses of both ankle flexor and extensor muscles are coaugmented in the early stance phase of human walking, suggesting that these coaugmented reflex responses contribute to secure foot stabilization around the heel strike. To test whether the reflex responses mediated by the stretch reflex pathway are actually induced in both the ankle flexor and extensor muscles when the supportive surface is suddenly destabilized, we investigated the electromyographic (EMG) responses induced after a sudden drop of the supportive surface at the early stance phase of human walking. While subjects walked on a walkway, the specially designed movable supportive surface was unexpectedly dropped 10 mm during the early stance phase. The results showed that short-latency reflex EMG responses after the impact of the drop (<50 ms) were consistently observed in both the ankle flexor and extensor muscles in the perturbed leg. Of particular interest was that a distinct response appeared in the tibialis anterior muscle, although this muscle showed little background EMG activity during the stance phase. These results indicated that the reflex activities in the ankle muscles certainly acted when the supportive surface was unexpectedly destabilized just after the heel strike during walking. These reflex responses were most probably mediated by the facilitated stretch reflex pathways of the ankle muscles at the early stance phase and were suggested to be relevant to secure stabilization around the ankle joint during human walking.     

Convergence in Reflex Pathways from Multiple Cutaneous Nerves Innervating the Foot Depends upon the Number of Rhythmically Active Limbs during Locomotion

Neural output from the locomotor system for each arm and leg influences the spinal motoneuronal pools directly and indirectly through interneuronal (IN) reflex networks. While well documented in other species, less is known about the functions and features of convergence in common IN reflex system from cutaneous afferents innervating different foot regions during remote arm and leg movement in humans. The purpose of the present study was to use spatial facilitation to examine possible convergence in common reflex pathways during rhythmic locomotor limb movements. Cutaneous reflexes were evoked in ipsilateral tibialis anterior muscle by stimulating (in random order) the sural nerve (SUR), the distal tibial nerve (TIB), and combined simultaneous stimulation of both nerves (TIB&SUR). Reflexes were evoked while participants performed rhythmic stepping and arm swinging movement with both arms and the leg contralateral to stimulation (ARM&LEG), with just arm movement (ARM) and with just contralateral leg movement (LEG). Stimulation intensities were just below threshold for evoking early latency (<80 ms to peak) reflexes. For each stimulus condition, rectified EMG signals were averaged while participants held static contractions in the stationary (stimulated) leg. During ARM&LEG movement, amplitudes of cutaneous reflexes evoked by combined TIB&SUR stimulation were significantly larger than simple mathematical summation of the amplitudes evoked by SUR or TIB alone. Interestingly, this extra facilitation seen during combined nerve stimulation was significantly reduced when performing ARM or LEG compared to ARM&LEG. We conclude that locomotor rhythmic limb movement induces excitation of common IN reflex pathways from cutaneous afferents innervating different foot regions. Importantly, activity in this pathway is most facilitated during ARM&LEG movement. These results suggest that transmission in IN reflex pathways is weighted according to the number of limbs directly engaged in human locomotor activity and underscores the importance of arm swing to support neuronal excitability in leg muscles.     

Interlimb Reflexes Induced by Electrical Stimulation of Cutaneous Nerves after Spinal Cord Injury

Whether interlimb reflexes emerge only after a severe insult to the human spinal cord is controversial. Here the aim was to examine interlimb reflexes at rest in participants with chronic (>1 year) spinal cord injury (SCI, n = 17) and able-bodied control participants (n = 5). Cutaneous reflexes were evoked by delivering up to 30 trains of stimuli to either the superficial peroneal nerve on the dorsum of the foot or the radial nerve at the wrist (5 pulses, 300 Hz, approximately every 30 s). Participants were instructed to relax the test muscles prior to the delivery of the stimuli. Electromyographic activity was recorded bilaterally in proximal and distal arm and leg muscles. Superficial peroneal nerve stimulation evoked interlimb reflexes in ipsilateral and contralateral arm and contralateral leg muscles of SCI and control participants. Radial nerve stimulation evoked interlimb reflexes in the ipsilateral leg and contralateral arm muscles of control and SCI participants but only contralateral leg muscles of control participants. Interlimb reflexes evoked by superficial peroneal nerve stimulation were longer in latency and duration, and larger in magnitude in SCI participants. Interlimb reflex properties were similar for both SCI and control groups for radial nerve stimulation. Ascending interlimb reflexes tended to occur with a higher incidence in participants with SCI, while descending interlimb reflexes occurred with a higher incidence in able-bodied participants. However, the overall incidence of interlimb reflexes in SCI and neurologically intact participants was similar which suggests that the neural circuitry underlying these reflexes does not necessarily develop after central nervous system injury.     

Stick insects walking on a wheel: Perturbations induced by obstruction of leg protraction

Summary Stick insects (Carausius morosus) walking on a wheel were perturbed by restricting the forward protraction of individual legs. A barrier placed before a single middle or rear leg prevented that leg from reaching its normal protraction endpoint but allowed it unimpeded retraction. Upon striking the barrier, the protracting leg attempted to get past it and thereby prolonged protraction. This prolongation increased with the extent to which the obstruction infringed upon the leg's normal step range. Barriers placed near the midpoint of this range elicited large perturbations: the blocked leg often continued its protraction throughout many step cycles of the other legs (Fig. 1 E, F). For the most part walking was irregular and smooth forward progression was disrupted. Nevertheless, the infrequent steps by the affected leg usually were coordinated with those of the adjacent ipsilateral legs.More rostral barrier positions elicited smaller perturbations: the blocked leg usually made one step in each step cycle of the other legs (Fig. 1 B, C, D, G). Measurements for these regular step sequences showed quantitatively that protraction duration increased in proportion to the severity of the infringement on normal leg movement (Figs. 3, 4). The fraction of the step period occupied by protraction increased from ca. 10% for normal walking to ca. 50% for caudal barrier positions. This proportionality is interpreted to show the importance of spatial components of the walking program.When one leg was obstructed, its extended protraction influenced the stepping of the three adjacent legs as follows. First, the ipsilateral rostral leg showed the largest change: its protraction onset was regularly delayed for the duration of the extended protraction (Figs. 4, 7, 8), demonstrating a strong, centrally mediated inhibition. The presence of a further delay of up to 100 to 140 ms suggests that peripheral input from the protracting leg may be important for releasing this inhibition. Second, steps by an adjacent caudal leg were not measurably affected. However, the method may not have sufficed to reveal such effects because during regular walking middle leg protractions rarely lasted long enough to conflict with subsequent steps by the ipsilateral rear leg. Third, contralateral effects differed between middle and rear leg obstructions. If the obstructed leg was a middle leg, its extended protraction had little effect upon stepping by the contralateral middle leg: the latter leg frequently protracted while the blocked leg continued its protraction and there was no consistent change in the phase relation of these two legs (Table 1). In contrast, if the obstructed leg was a rear leg, protractions by the contralateral rear leg tended to be delayed (Table 1).     

Three-dimensional modular control of human walking

Recent studies have suggested that complex muscle activity during walking may be controlled using a reduced neural control strategy organized around the co-excitation of multiple muscles, or modules. Previous computer simulation studies have shown that five modules satisfy the sagittal-plane biomechanical sub-tasks of 2D walking. The present study shows that a sixth module, which contributes primarily to mediolateral balance control and contralateral leg swing, is needed to satisfy the additional non-sagittal plane demands of 3D walking. Body support was provided by Module 1 (hip and knee extensors, hip abductors) in early stance and Module 2 (plantarflexors) in late stance. In early stance, forward propulsion was provided by Module 4 (hamstrings), but net braking occurred due to Modules 1 and 2. Forward propulsion was provided by Module 2 in late stance. Module 1 accelerated the body medially throughout stance, dominating the lateral acceleration in early stance provided by Modules 4 and 6 (adductor magnus) and in late stance by Module 2, except near toe-off. Modules 3 (ankle dorsiflexors, rectus femoris) and 5 (hip flexors and adductors except adductor magnus) accelerated the ipsilateral leg forward in early swing whereas Module 4 decelerated the ipsilateral leg prior to heel-strike. Finally, Modules 1, 4 and 6 accelerated the contralateral leg forward prior to and during contralateral swing. Since the modules were based on experimentally measured muscle activity, these results provide further evidence that a simple neural control strategy involving muscle activation modules organized around task-specific biomechanical functions may be used to control complex human movements.     

Responses of the lower limb to load carrying in walking man

Muscle activity patterns of several lower limb muscles were examined in the left leg of normal human subjects walking at comfortable speed on a treadmill. In addition knee angular changes and the durations of the swing and stance phases of the step cycle were recorded. Data were collected during a period of normal control walking and when the subject carried a load, either in his right or left hand or on his back. Load (up to 20% of body weight) carried in either hand caused minimal changes in the kinematic parameters investigated but evoked significant prolongation of the normal ongoing electromyographic activity in the contralateral Gluteus medius and in the ipsilateral Gastrocnemius, Vastus lateralis and Semimembranosus. Load (up to 50% of body weight) carried on the back significantly shortened the swing phase and prolonged the ongoing electromyographic activity of the Vastus lateralis. These findings would seem to indicate that the activity of the leg musculature during walking is so tightly controlled that deviation from the normal kinematic pattern of the legs is largely prevented even when body posture and balance are disturbed by carrying substantial additional load.     

Fusimotor reflexes influencing secondary muscle spindle afferents from flexor and extensor muscles in the hind limb of the cat.

The purpose of this study was to investigate secondary muscle spindle afferents from the triceps-plantaris (GS) and posterior biceps and semitendinosus (PBSt) muscles with respect to their fusimotor reflex control from different types of peripheral nerves and receptors. The activity of single secondary muscle spindle afferents was recorded from dissected and cut dorsal root filaments in alpha-chloralose anaesthetized cats. Both single spindle afferents and sets of simultaneously recorded units (2-3) were investigated. The modulation and mean rate of firing of the afferent response to sinusoidal stretching of the GS and PBSts muscle were determined. Control measurements were performed in the absence of any reflex stimulation, while test measurements were made during reflex stimulation. The reflex stimuli consisted of manually performed movements of the contralateral hind limb, muscle stretches, ligament tractions and electrical stimulations of cutaneous afferents. Altogether 21 secondary spindle afferents were investigated and 20 different reflex stimuli were employed. The general responsiveness (i.e. number of significant reflex effects/number of control-test series) was 52.4%, but a considerable variation between different stimuli was found, with the highest (89.9%) for contralateral whole limb extension and the lowest (25.0%) for stretch of the contralateral GS muscle. The size of the response to a given stimulus varied considerably between different afferents, and, in the same afferent, different reflex stimuli produced effects of varying size. Most responses were characterized by an increase in mean rate of discharge combined with a decrease in modulation, indicative of static fusimotor drive (Cussons et al., 1977). Since the secondary muscle spindle afferents are part of a positive feedback loop, projecting back to both static and dynamic fusimotor neurones (Appelberg Et al., 1892 a, 1983 b; Appelberg et al., 1986), it is suggested that the activity in the loop may work like an amplified which, during some circumstances, enhance the effect of other reflex inputs to the system (Johansson et al., 1991 b).     

A New Approach to the Analysis of Normal Muscle Electrical Activity during Walking in Humans

Introduction of short-term disturbances into the biomechanical structure of the human gait with simultaneous recording of the muscle electrical activity has been used to demonstrate that the human body has an intraspinal locomotor program consisting of an inhibition period, when afferent stimuli cause no response, and an excitation period, when the responses are expressed. The electromyographic profile of the leg muscles of subjects walking on a horizontal surface, up stairs, and down stairs may be divided into three zones. The H and M zones (the highest and moderate activities, respectively) correspond to the centrally programmed excitation period, and the L zone (low-amplitude activity), to the centrally programmed inhibition period. The difference between the former two zones is that the activity in the H zone is regular, whereas the M-zone activity is irregular and varies depending on biomechanical conditions. Apparently, the steady activity in the H zone is determined by the combined effect of the spinal generator of locomotion movements, cyclic supraspinal stimuli, and various (mainly proprioceptive) afferent impulses from the leg. An increase in the M-zone activity is mainly determined by afferent stimuli.     

Self-Sustained Motor Activity Triggered by Interlimb Reflexes in Chronic Spinal Cord Injury,Evidence of Functional Ascending Propriospinal Pathways

The loss or reduction of supraspinal inputs after spinal cord injury provides a unique opportunity to examine the plasticity of neural pathways within the spinal cord. In a series of nine experiments on a patient, quadriplegic due to spinal cord injury, we investigated interlimb reflexes and self-sustained activity in completely paralyzed and paretic muscles due to a disinhibited propriospinal pathway. Electrical stimuli were delivered over the left common peroneal nerve at the fibular head as single stimuli or in trains at 2–100 Hz lasting 1 s. Single stimuli produced a robust interlimb reflex twitch in the contralateral thumb at a mean latency 69 ms, but no activity in other muscles. With stimulus trains the thumb twitch occurred at variable subharmonics of the stimulus rate, and strong self-sustained activity developed in the contralateral wrist extensors, outlasting both the stimuli and the thumb reflex by up to 20 s. Similar behavior was recorded in the ipsilateral wrist extensors and quadriceps femoris of both legs, but not in the contralateral thenar or peroneal muscles. The patient could not terminate the self-sustained activity voluntarily, but it was abolished on the left by attempted contractions of the paralyzed thumb muscles of the right hand. These responses depend on the functional integrity of an ascending propriospinal pathway, and highlight the plasticity of spinal circuitry following spinal cord injury. They emphasize the potential for pathways below the level of injury to generate movement, and the role of self-sustained reflex activity in the sequelae of spinal cord injury.     

Intracellular recordings from nonspiking interneurons in a semiintact,tethered walking insect

Nonspiking interneurons were investigated in a tethered, walking insect, Carausius morosus, that was able to freely perform walking movements. Experiments were carried out with animals walking on a lightweight, double-wheel treadmill. Although the animal was opened dorsally, the walking system was left intact. Intracellular recordings were obtained from the dorsal posterior neuropil of the mesothoracic ganglion. Nonspiking inter-neurons, in which modulations of the membrane potential were correlated with the walking rhythm, were described physiologically and stained with Lucifer Yellow. Interneurons are demonstrated in which membrane potential oscillations mirror the leg position or show correlation with the motoneuronal activity of the protractor and retractor coxae muscles during walking. Other interneurons showed distinct hyperpolarizations at certain important trigger points in the step cycle, for example, at the extreme posterior position. Through electrical stimulation of single, nonspiking interneurons during walking, the motoneuronal activity in two antagonistic muscles—protractor and retractor coxae—could be reversed and even the movement of the ipsilateral leg could be influenced. The nonspiking interneurons described appear to be important premotor elements involved in walking. They receive, integrate, and process information from different leg proprioceptors and drive groups of leg motoneurons during walking.     

Cross-training effect of chronic whole-body vibration exercise: a randomized controlled study

Abstract

Purpose: To determine whether unilateral leg whole-body vibration (WBV) strength training induces strength gain in the untrained contralateral leg muscle. The secondary aim was to determine the potential role of spinal neurological mechanisms regarding the effect of WBV exercise on contralateral strength training.

Materials and Methods: Forty-two young adult healthy volunteers were randomized into two groups: WBV exercise and Sham control. An isometric semi-squat exercise during WBV was applied regularly through 20 sessions. WBV training was applied to the right leg in the WBV group and the left leg was isolated from vibration. Sham WBV was applied to the right leg of participants in the Control group. Pre- and post-training isokinetic torque and reflex latency of both quadricepses were evaluated.

Results: The increase in the strength of right (vibrated) knee extensors was 9.4?±?10.7% in the WBV group (p?=?.001) and was 1.2?±?6.6% in the Control group (p?=?.724). The left (non-vibrated) extensorsvibrated) knee extensors w4?±?8.4% in the WBV group (p?=?.038), whereas it decreased by 1.4?±?7.0% in the Control (p?=?.294). The strength gains were significant between the two groups. WBV induced the reflex response of the quadriceps muscle in the vibrated ipsilateral leg and also in the non-vibrated contralateral leg, though with a definite delay. The WBV-induced muscle reflex (WBV-IMR) latency was 22.5?±?7.7?ms for the vibrated leg and 39.3?±?14.6?ms for the non-vibrated leg.

Conclusions: Chronic WBV training has an effect of the cross-transfer of strength to contralateral homologous muscles. The WBV-induced muscular reflex may have a role in the mechanism of cross-transfer strength.     

Modulation of segmental reflex reactions during actual locomotion in rats

High- and low-threshold reflex segmental reflex reactions produced by stimulating the dorsal root at different stages of the locomotor cycle were investigated during locomotor swimming motions in white rats. Findings show respectively considerable inhibition of and a pronounced increase in extensor and flexor lowthreshold reflex reactions during the absence and presence of activity in the associated muscles. Low-threshold stimulation produced no outstanding effect on the shaping of the muscles' own activity and hence failed to affect time course or amplitude parameters of locomotor movements. Changes in reflex reactions to high-threshold stimulation during the locomotor cycle largely resembled changes in these responses to low-threshold stimulation, although development of highthreshold reactions differed from that of low-threshold response in affecting the parameters of locomotor activity in the associated muscle, while likewise altering frequency parameters of the locomotor rhythm. The physiological significance and mechanisms possibly underlying modulations in the efficacy of afferent peripheral influences during locomotion are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 3, pp. 326–333, May–June, 1988.     

Intracellular recordings from nonspiking interneurons in a semiintact, tethered walking insect.

Nonspiking interneurons were investigated in a tethered, walking insect, Carausius morosus, that was able to freely perform walking movements. Experiments were carried out with animals walking on a lightweight, double-wheel treadmill. Although the animal was opened dorsally, the walking system was left intact. Intracellular recordings were obtained from the dorsal posterior neuropil of the mesothoracic ganglion. Nonspiking interneurons, in which modulations of the membrane potential were correlated with the walking rhythm, were described physiologically and stained with Lucifer Yellow. Interneurons are demonstrated in which membrane potential oscillations mirror the leg position or show correlation with the motoneuronal activity of the protractor and retractor coxae muscles during walking. Other interneurons showed distinct hyperpolarizations at certain important trigger points in the step cycle, for example, at the extreme posterior position. Through electrical stimulation of single, nonspiking interneurons during walking, the motoneuronal activity in two antagonistic muscles--protractor and retractor coxae--could be reversed and even the movement of the ipsilateral leg could be influenced. The nonspiking interneurons described appear to be important premotor elements involved in walking. They receive, integrate, and process information from different leg proprioceptors and drive groups of leg motoneurons during walking.     

Mucosal afferents mediate laryngeal adductor responses in the cat.

Laryngeal adductor responses (LAR) close the airway in response to stimulation of peripheral afferents in the superior laryngeal nerve. Although both mucosal afferents and proprioceptive receptors are present in the larynx, their relative contribution for reflex elicitation is unknown. Our purpose was to determine which receptor types are of importance in eliciting the LAR. A servomotor with displacement feedback was used to deliver punctate displacements to the body of the arytenoid cartilage and overlying mucosa on each side of the larynx in eight anesthetized cats. The same displacements were delivered both before and after surgical excision of the overlying mucosa. With the mucosa intact, early short-latency component R1 LAR responses recorded from the thyroarytenoid muscles were frequent (ipsilateral > 92%, contralateral > 95%). After the mucosa was removed, the LAR became infrequent (<3%) and was reduced in amplitude in both the ipsilateral and contralateral thyroarytenoid muscle recording sites (P < 0.0005). These findings demonstrate that mucosal mechanoreceptors and not proprioceptive afferents contribute to the elicitation of LAR responses in the cat.     

Homologous Muscle Contraction during Unilateral Movement Does Not Show a Dominant Effect on Leg Representation of the Ipsilateral Primary Motor Cortex

Co-activation of homo- and heterotopic representations in the primary motor cortex (M1) ipsilateral to a unilateral motor task has been observed in neuroimaging studies. Further analysis showed that the ipsilateral M1 is involved in motor execution along with the contralateral M1 in humans. Additionally, transcranial magnetic stimulation (TMS) studies have revealed that the size of the co-activation in the ipsilateral M1 has a muscle-dominant effect in the upper limbs, with a prominent decline of inhibition within the ipsilateral M1 occurring when a homologous muscle contracts. However, the homologous muscle-dominant effect in the ipsilateral M1 is less clear in the lower limbs. The present study investigates the response of corticospinal output and intracortical inhibition in the leg representation of the ipsilateral M1 during a unilateral motor task, with homo- or heterogeneous muscles. We assessed functional changes within the ipsilateral M1 and in corticospinal outputs associated with different contracting muscles in 15 right-handed healthy subjects. Motor tasks were performed with the right-side limb, including movements of the upper and lower limbs. TMS paradigms were measured, consisting of short-interval intracortical inhibition (SICI) and recruitment curves (RCs) of motor evoked potentials (MEPs) in the right M1, and responses were recorded from the left rectus femoris (RF) and left tibialis anterior (TA) muscles. TMS results showed that significant declines in SICI and prominent increases in MEPs of the left TA and left RF during unilateral movements. Cortical activations were associated with the muscles contracting during the movements. The present data demonstrate that activation of the ipsilateral M1 on leg representation could be increased during unilateral movement. However, no homologous muscle-dominant effect was evident in the leg muscles. The results may reflect that functional coupling of bilateral leg muscles is a reciprocal movement.