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.     

The patterns of changes in the electromyographic profile of muscles in some gait disorders

Electromyograms (EMGs) of subjects with different gait pathologies have been used to analyze the most typical forms of transformation of the muscle EMG profile: (1) a decrease in the muscle electrical activity during a locomotor cycle, either uniform or with a predominant reduction in the region of maximum values (the Mx zone); the former variant is characteristic of structural damage to nerves and muscles, whereas the latter variant is of functional origin because it is related to the alterations in the biomechanical conditions of muscle work during walking; (2) an increase in muscle electrical activity during the cycle, its maxima being prolonged from the Mx zone to the zone of moderate activity (the Md zone); and (3) a complete shift of the activity peaks to another phase of the cycle, usually from the Mx to the Md zone. It has been assumed that all forms of the transformation of the muscle EMG profile except a drastic uniform decrease in activity on damage to nerve and muscle structures are of afferent origin. These types of alterations in the EMG profile are accounted for by disturbance of the biomechanical conditions of muscle activity, such as an increased or decreased muscle load.     

Active selection of afferent information as the basic principle of peripheral correction of the activity of spinal rhythmic movement generators

This paper summarizes information obtained in the experimental study of the dynamics of polarization of central primary afferent endings and modifications of segmental responses to afferent stimuli during fictitious locomotion and fictitious scratching in immobilized, decorticated, decerebrate, and spinal cats. Fictitious locomotion was accompanied by tonic hyperpolarization, fictitious scratching by tonic depolarization of central primary afferent endings. Against the background of these long-lasting changes in primary afferent depolarization, it exhibited periodic changes in the rhythm of efferent activity. Periodic changes of depolarization were virtually in phase in different ipsilateral segments of the lumbosacral enlargement. Data on groups of afferent fibers in whose central endings tonic and phasic changes of polarization took place. The appearance of fictitious locomotion was accompanied by a tonic increase, and of fictitious scratching by tonic inhibition of several evoked segmental responses. These tonic changes were a background against which segmental responses were modulated in step with the working rhythm of the locomotion and scratching generators. Many of the changes in evoked segmental responses were shown to be based on modulation of polarization of central endings of primary afferents by locomotion and scratching generators. It is concluded that active tonic and phase-dependent selection of incoming afferent information is effected through modulation of presynaptic inhibition of the generator. The role of this selection in peripheral collection of activity of locomotion and scratching generators is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 343–353, May–June, 1984.     

Age-related parameters of spinal inhibition of skeletal muscles in regulation of voluntary movements in men

Age-related features of spinal inhibition in the regulation of voluntary movements in men were studied. It was found that presynaptic and nonreciprocal and reciprocal inhibitions of the flexor of toes during voluntary movements were less intense than inhibition during relative muscle rest in subjects of all age groups. This results from the age-related features of supraspinal excitatory and inhibitory effects on Ia and Ib spinal interneurons, which change the mechanism of spinal inhibition of voluntary movements as the organism develops. In boys 9–12 years of age, execution of voluntary movements is accompanied by the lowest presynaptic inhibition of Ia afferents and the most pronounced increase in nonreciprocal and reciprocal inhibition of α-motoneurons in the flexor of toes, compared to the other age groups. Execution of voluntary movements by boys 14–15 years of age leads to an increase in presynaptic inhibition of Ia afferents and the most pronounced decrease in reciprocal and nonreciprocal inhibition of spinal α-motoneurons of the flexor of toes. By the age of 17–18 years, the mechanism of nonreciprocal inhibition of α-motoneurons of the flexor of toes during voluntary movements is similar to that in adolescents aged 14–15 years. The definitive level of presynaptic inhibition of Ia afferents and reciprocal inhibition of α-motoneurons of the flexor of toes during voluntary movements is reached by 17–18 years.     

Kinematics of different types of locomotor movements in the deafferented rat

The kinematics of rat hindlimb movements were assessed and compared pre- and post-deafferentation during swimming, forelimb treadmill locomotion plus hindlimb swimming motion, and walking using all four limbs. All types of locomotion were characterized by an increase in the frequency of locomotor rhythm and reduced amplitude of motion at the hindlimb joints following deafferentation. The reduced change observed in the angle of the coxofemoral joint, indicative of a horizontal component in locomotor motion, was mainly brought about by less marked extension. This would confirm evidence indicating that increased load on the extremities, with its ensuing naturally-occurring afferent outflow, is accompanied by a reduced locomotor motion rate and a rise in the amplitude of the latter due to intensified extension of the limb. The increased forward carriage of the hind limb seen during the transition to four-legged locomotion persisted after deafferentation; this may be considered a sign of coordination amongst the limbs. Deafferentation led to a reduction in the MEG of muscle activity, which was found to be lowest in swimming and highest during walking. The role of the afferent inflow in shaping different types of locomotor motion is evaluated.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 19, No. 4, pp. 520–525, July–August, 1987.     

Synaptic and electotonic contacts on primary afferent axons in the lamprey <Emphasis Type="Italic">Lampetra fluviatilis</Emphasis> spinal cord

Distribution of GABA and glycine immunoreactivity was studied in synapses on primary afferent axons of the lamprey Lampetra fluviatilis spinal cord using a double labelling technique. Approximately 25% of synapses exhibit GABA immunoreactivity, while more than 70% are immunoreactive to both neurotransmitters. As in other vertebrates, axo-axonal contacts represent three-component synaptic complexes, the so-called triads, where the immunoreactive terminal make synaptic contact simultaneously with the afferent axon and the dendrite contacting this afferent. Contact zones with gap junction-like cell membrane specializations were found between adjacent afferents suggesting the presence of electrotonic interaction between them. This interaction appears to serve for the synchronization of the afferent flow and represents a structural correlate of the mechanism of rapid interneuronal communication between functionally uniform neurons, which is an important element in the organization of coordinated locomotor acts. Besides, our studies provide evidence that afferent–afferent interaction may be mediated not only electrotonically but also with the aid of chemical synapses. This finding suggests that glutamate-induced depolarization of primary afferents results not only from autoreception but also from the direct effect of glutamate on the afferent’s cell membrane.     

Somatosensory unit input to the spinal cord during normal walking

Chronic recording techniques in freely walking cats have been used to sample unitary activity from most large myelinated afferent classes. Cutaneous mechanoreceptors are highly sensitive and generate regular activity patterns predictable from their modalities. Knee joint afferents can fire briskly midrange locomotory movements but appear to be influenced by factors other than joint angle. Golgi tendon organs generate activity consistent with sensitivity to active muscle tension. Muscle spindle afferents do not appear to conform to any single functional pattern for all muscles. It is suggested that degree and rate of stretch are sensed by spindles (possibly under dynamic fusimotor bias) in extensor muscles which normally undergo isometric or lengthening contractions whereas rapidly modulated static fusimotor activity is employed to preserve spindle activity during the rapidly shortening contractions of flexor muscles. Both patterns may be represented in different spindles of bifunctional, biarticular muscles such as rectus femoris and sartorius.     

Phase-dependent changes in dorsal root potential during actual locomotion in rats

Fluctuations in dorsal root potential (DRP) were investigated in trials on white rats during two types of locomotion, differing in the intensity of afferent flow (swimming and walking). Two negative waves of DRP were observed corresponding to the stance (or propulsive) phase and the swing (or transfer) phase within a single locomotor cycle. Whereas DRP had risen primarily during the stroke phase with increased intensity during swimming, it increased during the standing phase in walking. A relationship was revealed between the amplitude of DRP and the intensity of afferent flow apparent during passive displacement of the limb, as well as locomotion. It is concluded that DRP waves are mainly due to influences from peripheral afferents during actual locomotion.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 3, pp. 333–340, May–June, 1988.     

Limb movement,intensity of integral afferent impulses from limb receptors,and level of primary afferent terminal polarization during locomotion in cats decerebrated at high level

The relationship between parameters of electrical muscle activity, changes at hindlimb joint angles, intensity of integral afferent flow, and dorsal root potential during real-life locomotion was investigated in cats decerebrated at high level. Characteristics of rear limb movements before and after deafferentation were described. It was found that afferent activity during locomotion motion consists, of two components — a tonic and a periodic phasic stage. Three main waves may be distinguished in the latter, each of which gives rise to associated changes in the level of primary afferent terminal polarization. These changes in turn are summated with the effects produced by the central generator. Correlations, between the parameters of these processes were investigated and the mechanisms underlying afferent control of locomotion generator function discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 1, pp. 119–127, January–February, 1988.     

Immunoreactivity of synapses on primary afferent axons and sensory neurons in lamprey spinal cord (<Emphasis Type="Italic">Lampetra fluviatilis</Emphasis>)

The ultrastructure and immunospecificity of synapses on primary afferents and dorsal sensory cells (DCs) were studied in lamprey (Lampetra fluviatilis) spinal cords. Using the postembedding immunogold method with a combination of antibodies—polyclonal antibodies to glutamate and monoclonal antibodies to gamma-aminobutyric acid (GABA)—the presence of GABA-positive on the primary afferent axons and GABA-and glutamate-immunopositive synapses on the DC somatic membranes have been shown. Thus, it is obvious that sensory information in the lamprey is controlled by both presynaptic inhibition via synapses on the primary afferent axons and by direct synaptic influence on the body of the sensory neuron.     

Comparison of Trunk Activity during Gait Initiation and Walking in Humans

To understand the role of trunk muscles in maintenance of dynamic postural equilibrium we investigate trunk movements during gait initiation and walking, performing trunk kinematics analysis, Erector spinae muscle (ES) recordings and dynamic analysis. ES muscle expressed a metachronal descending pattern of activity during walking and gait initiation. In the frontal and horizontal planes, lateroflexion and rotation occur before in the upper trunk and after in the lower trunk. Comparison of ES muscle EMGs and trunk kinematics showed that trunk muscle activity precedes corresponding kinematics activity, indicating that the ES drive trunk movement during locomotion and thereby allowing a better pelvis mobilization. EMG data showed that ES activity anticipates propulsive phases in walking with a repetitive pattern, suggesting a programmed control by a central pattern generator. Our findings also suggest that the programs for gait initiation and walking overlap with the latter beginning before the first has ended.     

Isozyme Analysis in a Genetic Collection of Amaranths (<Emphasis Type="Italic">Amaranthus</Emphasis> L.)

In various populations of the cultivated and weedy amaranth species, the electrophoretic patterns of alcohol dehydrogenase (ADH), glutamate dehydrogenase (GDH), malate dehydrogenase (MDH), isocitrate dehydrogenase (IDH) and malic enzyme (Me) were studied. In total, 52 populations and two varieties (Cherginskii and Valentina) have been examined. Allozyme variation of this material was low. Irrespective of species affiliation, 26 populations and two varieties were monomorphic for five enzymes; a slight polymorphism of three, two, and one enzymes was revealed in three, nine, and fourteen populations, respectively. A single amaranth locus, Adh, with two alleles, Adh F and Adh S, controls amaranth ADH. Two alleles, common Gdh S and rare Gdh F, control GDH; no heterozygotes at this locus were found. The MDH pattern has two, the fast- and slow-migrating, zones of activity (I and II, respectively). Under the given electrophoresis conditions, the fast zone is diffuse, whereas slow zone is controlled by two nonallelic genes, monomorphic Mdh 1 and polymorphic Mdh 2 that includes three alleles: Mdh 2-F, Mdh 2-N, and Mdh 2-S. Low polymorphism of IDH and Me was also found, though their genetic control remains unknown.     

Calcitonin gene-related peptide and capsaicin inhibit the circular muscle of the guinea-pig ileum

Calcitonin gene-related peptide (CGRP) is known to excite, through the release of acetylcholine, the circular muscle (CM) of the guinea-pig ileum in vitro. In the present experiments, the effect of rat CGRP was investigated on the CM of tetrodotoxin-treated, spontaneously active ileum preparations. CGRP (1-10 nM) caused concentration-dependent inhibition of both the amplitude and frequency of spontaneous CM contractions. Capsaicin (a sensory stimulant known to release CGRP from primary afferents) also inhibited CM activity. The effect of 1 microM capsaicin underwent rapid desensitization, indicating specific action on afferent structures, whereas a high concentration of the drug (33 microM) inhibited CM activity most probably on the smooth muscle itself.     

The effects of sloped surfaces on locomotion: backward walking as a perturbation

The purpose of this study was to examine lower extremity kinetics and muscle activity during backward slope walking to clarify the relationship between joint moments and powers and muscle activity patterns observed in forward slope walking. Nine healthy volunteers walked backward on an instrumented ramp at three grades (-39% (-21 degrees ), 0% (level), +39% (+21 degrees )). EMG activity was recorded from major lower extremity muscles. Joint kinetics were obtained from kinematic and force platform data. The knee joint moment and power generation increased significantly during upslope walking; hip joint moment and power absorption increased significantly during downslope walking. When compared to data from forward slope walking, these backward walking data suggest that power requirements of a task dictate the muscle activity pattern needed to accomplish that movement. During downslope walking tasks, power absorption increased and changes in muscle activity patterns were directly related to the changes in the joint moment patterns. In contrast, during upslope walking tasks, power generation increased and changes in the muscle activity were related to the changes in the joint moments only at the 'primary' joint; at adjacent joints the changes in muscle activity were unrelated to the joint moment pattern. The 'paradoxical' changes in the muscle activity at the adjacent joints are possibly related to the activation of biarticular muscles required by the increased power generation at the primary joint. In total, these data suggest that changing power requirements at a joint impact the control of muscle activity at that and adjacent joints.     

Modulation of limb dynamics in the swing phase of locomotion

A method was presented for quantifying cat (Felis catus) hind limb dynamics during swing phase of locomotion using a two-link rigid body model of leg and paw, which highlighted the dynamic interactions between segments. Comprehensive determination was made of cat segment parameters necessary for dynamic analysis, and regression equations were formulated to predict the inertial parameters of any comparable cat. Modulations in muscle and non-muscle components of knee and ankle joint moments were examined at two treadmill speeds using three gaits: (a) pace-like walk and trot-like walk, at 1.0 ms-1, and (b) gallop, at 2.1 ms-1. Results showed that muscle and segment interactive moments significantly effected limb trajectories during swing. Some moment components were greater in galloping than in walking, but net joint maxima were not significantly different between speeds. Moment magnitudes typically were greater for pace-like walking than for trot-like walking at the same speed. Generally, across gaits, the net and muscle moments were in phase with the direction of distal joint motion, and these same moments were out of phase with proximal joint motion. Intersegmental dynamics were not modulated exclusively by speed of locomotion, but interactive moments were also influenced significantly by gait mode.     

Cephalic anatomy and three-dimensional reconstruction of the head of <Emphasis Type="Italic">Catops ventricosus</Emphasis> (Weise, 1877) (Coleoptera: Leiodidae: Cholevinae)

Adult head structures are well known in the coleopteran suborders Archostemata and Adephaga, whereas the available information is very fragmentary in the megadiverse Polyphaga, including the successful superfamily Staphylinoidea. In the present study, the cephalic morphology of the cholevine species Catops ventricosus is described in detail and documented. The results were compared to conditions occurring in other polyphagan lineages, especially staphylinoid and scarabaeoid representatives. Specific external features documented in Catops and potential autapomorphies of Leiodidae include a five-segmented antennal club with a reduced eighth antennomere and the presence of periarticular grooves filled with sensilla on antennomeres 7, 9, and 10. The firm connection of the head and pronotum is possibly an apomorphy of Cholevinae. The monophyly of Cholevinae excluding Eucatopini and Oritocatopini is supported by the apical maxillary palpomere as long as or shorter than the subapical one, and the presence of cryptic pore plates on the surface of these palpomeres—a feature described and documented here for the first time. The internal cephalic structures of Catops are mostly plesiomorphic, as for instance the complete tentorium. The pattern of the muscles is similar to what is found in other staphylinoid taxa. The unusual maxillary muscle “Mx” is likely a groundplan apomorphy of the clade Staphyliniformia?+?Scarabaeoidea. M. hypopharyngomandibularis (M13) was identified in Catops and is ancestral for Coleoptera, even though it is often missing. The same applies to M. tentoriohypopharyngalis (M42).     

Projections of Olfactory Bulbs and Non-Olfactory Telencephalic Structures in Amygdaloid Complex of the Turtle <Emphasis Type="Italic">Testudo horsfieldi</Emphasis>: A Study Using Anterograde Tracer Technique

To confirm the discrete character of projections of telencephalic olfactory and non-olfactory structures to the amygdaloid complex (AC) in the terrestrial turtle Testudo horsfieldi, a study was performed by the method of anterograde axonal transport of tracers (HRP, BDA). After a massive injection of the tracers into the main and accessory olfactory bulb, a dense accumulation of labeled fibers and terminals was found in ventral part of AC in the neuropil zones of nbam (J) and ncoam and very scanty—in nmam and ncam. After a massive injection of the tracers into non-olfactory telencephalic structures including dorsal cortex, pallidal enlargement, and ADVR, a very dense terminal field was observed in the dorsal AC part and a less dense one, with predominance of labeled fibers, in its ventral part. Local administration of the tracers separately into the dorsolateral (visual area) and the ventromedial (auditory-somatic area) parts of the ADVR allowed revealing discrete projections, respectively, to the laterocentral and mediocentral areas of the dorsal AC part with a relative overlapping in the central AC area. In all experiments, retrogradely labeled neurons in AC were also observed in zones of the corresponding bulbar and rostrotelencephalic projections. Thus, it has been shown that in the turtle AC there exist not only separation of direct olfactory and non-olfactory projections, but also discrete projections of different sensory areas of ADVR. Reciprocity of these connections is also confirmed. Organization of afferent olfactory and non-olfactory telencephalic connections in AC is similar in reptiles and in mammals.     

Recognition of fragmented images and mechanisms of memory

Analysis of the topography and parameters of event-related potentials (ERPs) recorded during the presentation of incomplete images with different fragmentation aided in study of the role of different cortical zones and the order of their involvement in the recognition process. The role of the frontal cortical areas at different stages of perception of fragmented images was established. The differences in the ERPs induced by recognized and unrecognized stimuli in the frontal and frontal-temporal derivations in the interval 30–83 ms were associated with the appearance of early positivity in response to recognized images and development of early negativity in response to unrecognized stimuli. The N300 component associated with recognition was stronger in these cortical zones during identification of images. A late positive complex appeared in the frontal areas earlier than in other areas. Involvement of the caudal visual areas in the recognition process was reflected by enhancement of the components P100, P250, and N400. Our results suggest that the frontal areas play the main role in the recognition of fragmented images because they are the structures that organize extraction of traces from long-term modality-specific memory using a system of afferent and efferent links and determine the strategy of information analysis necessary for the solution of a given task.