Responses of muscle spindles of tenotomized and hypertrophied muscles to stretching and vibration

Responses of muscle spindles of tenotomized and hypertrophied muscles to stretching and vibration were investigated. During constant stretching of the muscles with a load of 100 g the spontaneous activity of the primary endings in the control muscle was 17±1.5 spikes/sec, in the hypertrophied muscle it was unchanged, and after tenotomy it was increased to 26±1.5 spikes/sec. The discharge frequency of the secondary endings was unchanged under these circumstances. Responses of primary and secondary endings of spindles of the tenotomized muscle during the dynamic and static phases of stretching were higher in frequency than responses of spindles of normal muscles. The discharge frequency of the primary endings in the hypertrophied muscle also was increased during both phases of stretching. Responses of secondary endings of the spindles of the hypertrophied muscle were indistinguishable under these circumstances from responses of normal muscles. Primary endings of spindles of tenotomized and hypertrophied muscles, just as normally, reproduced frequencies of vibration stimulation up to 2000 Hz, but some increase in the discharge frequency was observed in the secondary endings at this time.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 8, No. 3, pp. 311–317, September, 1976.     

The relative sensitivity of Renshaw cells to orthodromic group Ia volleys caused by static stretch and vibrations of extensor muscles.

1. Activity of Renshaw cells monosynaptically excited by ventral root stimulation and disynaptically excited by electric stimulation of the group Ia afferents in the gastrocnemius-soleus (GS) nerve, was recorded in precollicular decerebrate cats. The response of these units to prolonged vibration applied longitudinally to the deefferented GS muscle was then compared with that elicited by static stretch of the homonymous muscle, for comparable frequencies of discharge of the group Ia afferents. 2. Small-amplitude vibration of the GS muscle at 200/sec for one second produced a sudden increase in the discharge rate of Renshaw cells, which gradually decreased within the first 100 msec of vibration to reach steady albeit lower level than that obtained during the first part of vibration. The response of the Renshaw cells during the first 100 msec of vibration (phasic response) and that elicited during the last 500 msec of vibration (tonic response) were evaluated for different frequencies of sinusoidal stretch. The mean increase in the firing frequency per imp./sec in the Ia afferents was also calculated using the total one-second period. 3. The response of Renshaw cells to muscle vibration increased with the frequency of vibration and, over the value of 10/sec, appeared to be linearly related to the frequency of the input, at least up to the frequency of 150/sec. Since vibration was of sufficient amplitude to produce driving of all the primary endings of muscle spindles, the responses were expressed as mean increases in the discharge rate of Renshaw cells per average impulse/sec in the Ia afferents. The discharge of the Renshaw cell increased on the average by 2.90 and 1.08 imp./sec per each imp./sec in the Ia afferents during the phasic and the tonic component of the response respectively, while the response calculated during the whole period of vibration corresponded on the average to 1.45 imp./sec per each imp./sec in the Ia afferents. 4. The Renshaw cells tested above responded also with increasing frequencies of discharge to increasing levels of static extension of the GS muscle. In particular the discharge frequency of Renshaw cells was on the average linearly related to muscle extension, at least for values ranging from 0 to 8 mm. The mean increase in discharge rate as a function of the static extension corresponded on the average to 0.89 imp./sec/mm. Since the discharge rate of the primary endings of muscle spindles recorded from the deefferented GS muscle increased by 2.62 imp./sec/mm, it appears that the mean increase in the discharge rate of Renshaw cells as a function of static extension corresponded to 0.34 imp./sec per each imp./sec in the Ia afferents.     

Types of spontaneous discharges of muscle receptors

Characteristics of discharges of 462 muslce spindle receptors located in the deefferented triceps surae muscle with a divided tendon were analyzed in anesthetized cats. Of the total number, 205 units had spontaneous activity which, depending on its stability, could be divided into three types. A long continuous spike train was generated by 87% of units. Distributions of interspike intervals were unimodal and symmetrical. Spontaneous activity of this type was more regular in secondary than in primary endings. Action potentials in 8% of units were grouped in volleys and interval distributions were bimodal. Spontaneous activity of the remaining 5% of units was characterized by sporadically appearing spikes with long intervals between them. The first two types of spontaneous activity cannot be placed in the category of stochastic processes. Threshold discharges during static stretching of the muscle sufficient to cause the appearance of a long spike train were studied in 231 of 257 initially "silent" units. By the same criteria these discharges were divided into corresponding types. Comparison of the characteristics of spontaneous activity and evoked responses suggests that they share the same mechanism of origin. It is concluded that the spontaneous discharge can be regarded as a static response of the receptors due not to an externally applied stretching load, but to the action of internal factors, depending on interaction between extrafusal and intrafusal muscle fibers.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 315–321, May–June, 1981.     

Spontaneous unit activity in anterior (limbic) nuclei of rabbit thalamus

Spontaneous spike activity in three anterior (limbic) neurons of the thalamic nucleus was studied by means of extracellular recording during chronic experiments on anesthetized rabbits. Neurons of the anteroventral nucleus showed high mean rate (24.8±5.8 spikes/sec) and varying structure of spike discharges ("inactivating" bursts of discharges, modulations in delta- and theta-rhythms, and bursts of discharges with a spindle rhythm of 12–14 Hz). "Inactivating" bursts of discharges alternating with single discharges predominated in the activity of neurons of the anteromedial nucleus (mean rate 10.0±1.4 spikes/sec). Activity of the anterodorsal nucleus could be clearly distinguished by the predominance of high-frequency groups of spikes (mean group frequency 67±5 spikes/sec) with prolonged intervals between groups.Institute of Biological Physics, Academy of Sciences of the USSR, Puschino, Moscow Province. Translated from Neirofiziologiya, Vol. 17, No. 5, pp. 579–586, September–October, 1985.     

Responses of muscle spindles of fast and slow muscles to mechanical stimulation during hypokinesia

Activity of muscle spindles of fast (extensor digitorum longus) and slow (soleus) muscles was studied in cats during hypokinesia of the limb immobilized in a plaster cast. Spontaneous activity of muscle spindles of the fast and slow muscles was unchanged during hypokinesia. Spontaneous activity of primary and secondary endings evoked by passive stretching of the muscle exceeded normal. During stretching of the muscles at different speeds and of different amplitudes, the discharge frequency of the primary and secondary endings was much greater than normally during both the dynamic and the static phase of stretching. These changes were more marked in the slow muscles.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 2, pp. 186–192, March–April, 1978.     

Spontaneous activity of passive muscle spindles of the cat triceps surae muscle

The hypothesis that one possible cause of the spontaneous discharge of muscle spindles in the totally relaxed and de-efferented triceps surae muscle, with its tendon divided, is the mechanical factor due to extrafusal-intrafusal interaction was tested in experiments on anesthetized cats. Responses of spontaneously active units were similar with respect to many indices to responses of silent receptors capable of generating a long discharge in response to an increase in static length of the muscle. Isotonic contraction of the relaxed muscle during direct or indirect stimulation was accompanied by a pause in spontaneous activity. The prolonged increase in discharge frequency sometimes arising as a result of pressure on the muscle in the region of the receptor also was abolished by weak isotonic contraction of the muscle. If a long posttetanic sensory discharge was induced in a spontaneously active receptor by intensive tetanization of the nerve to the muscle, and it was then abolished by short stretching of the muscle, the initial spontaneous discharge frequency was restored. The dynamic thresholds of spontaneously active receptors were lower than for silent receptors. In some spontaneously active receptors an initial slowing of the discharge was observed during linear or stepwise stretching of the muscle. It is suggested that the ability of sensory endings to generate a long spontaneous discharge is due to initial stretching of the spindle inside the relaxed muscle.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 3, pp. 287–294, May–June, 1978.     

Response of Renshaw cells to sinusoidal stretch of hindlimb extensor muscles.

1. Renshaw cells responding disynaptically to electrically induced group I volleys in the intact gastrocnemius-soleus (GS) nerve, were submitted to small-amplitude, high-frequency vibration applied longitudinally to the deefferented GS muscle in precollicular decerebrate cats. 2. Vibration of the GS muscle at 200/sec, 180 mu peak-to-peak amplitude for 80-100 msec produced a sudden increase in the discharge rate of Renshaw cells, which gradually decreased within 25-50 msec to reach a steady level higher than that recorded in the absence of vibration. 3. Excitation of Renshaw cells appeared at a threshold amplitude of vibration (at 200-250/sec) of 5-20 mu and increased to a maximum value for amplitudes of about 70-80 mu, i.e., when all the primary endings of the spindles from the GS muscle had been driven by the stimulus. Recruitment of the secondary endings of the muscle spindles, due to large amplitude muscle vibration, did not modify the response of the Renshaw cells to the mechanically induced group Ia volleys. 4. These findings were obtained with the GS muscle pulled at 8 mm of initial extension. A threshold response of Renshaw cells to vibration appeared at 4 mm of static stretch, while maximal responses occurred at 8 mm. No further increase and actually a slight decrease in the response appeared for initial extensions of the muscle of 10-12 mm. 5. For a given vibration amplitude, the response of the Renshaw cells increased with increasing frequencies of vibration to reach the maximum at frequencies of 150-250/sec. Bursts of Renshaw cell discharges synchronous to each stroke of vibrator occurred only for low frequencies of stimulation (less than 25/sec). 6. It is concluded that vibration of the GS muscle represents a very effective method in exciting the Renshaw cells and that this response depends upon selective stimulation of homonymous motoneurons monosynaptically excited by the orthodromic volleys originating from the primary endings of the corresponding muscle spindles.     

Tranformation of long reticulofugal spike trains by interneurons connected monosynaptically with the reticulospinal tract

Characteristics of transmission of activity evoked by stimulation of the reticular formation through interneurons located in the ventromedial zones of the gray matter of the lumbar division of the spinal cord and connected monosynaptically with reticulospinal fibers were investigated in cats. Responses of the neurons to relatively low-frequency (not exceeding 80–100/sec) stimulation consisted mainly of stationary discharges; with a further increase in frequency the response became nonstationary (the initial, relatively high-frequency discharge was followed by partial or complete suppression of the discharge). The maximal frequency of the initial phase of the response to high (over 400/sec) frequencies of stimulation was 180–230 spikes/sec. The "transmission factor" (ratio between the frequency of spikes in the response to the frequency of stimulation), calculated for stationary discharges, reached 0.7–0.8 at low frequencies of stimulation, and then decreased significantly. On the basis of the statistical characteristics of the stationary portions of the evoked activity and analysis of these data by the use of a mathematical model, indirect estimates were obtained of the parameters of processes lying at the basis of the transforming properties of this cell population.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 3, pp. 278–286, May–June, 1978.     

Types of activity of skeletal muscle motor units during cold tremor in cats

Discharges of 184 motor units of the sartorius muscle functioning during cold tremor were investigated in acute experiments on anesthetized cats. Units whose discharges correlated with respiration cycles and units functionally independent of the rhythm of respiration were discovered. Discharges of both types of motor units possessed the same mean frequency (4–12 spikes/sec) and the same low variability of interspike intervals. Additional temperature stimulation of the vascular temperature receptors and changes in the frequency and depth of the respiratory excursions during cold tremor evoked identical responses in the two types of units. It is concluded that both types of motor units have similar thresholds and are of the slow, phasic type.O. V. Kuusinen State University, Petrozavodsk. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 355–361, July–August, 1979.     

Time coupling of skeletomotor discharges in response to pseudo-random transsynaptic and transmembrane stimulation

Firing pattern of skeletomotor neurones innervating triceps surae muscles in response to pseudorandom muscle stretching and white noise modulated transmembrane current stimulation was investigated in decerebrate cats. Pseudo-random muscle stretching (upper cut-off frequency 60 Hz, amplitude (standard deviation) ranging from 18.5 m to 40 m) was applied to triceps surae muscles. Membrane potential changes and action potentials of skeletomotor neurones were recorded intracellularly. White noise modulated current was applied through the same (recording) microelectrode. Sequences of ten identical 5 s periods of either muscle stretching or transmembrane current stimulation were applied. Skeletomotor neurones belonging to slow motor units (rheobase less than 8.5 nA) generated action potentials in response to both pseudo-random muscle stretching and transmembrane current stimulation, while firing threshold of those belonging to fast motor units could not be reached by the muscle stretches applied. Peri-spike averaging of muscle length and injected current records showed that the action potentials appeared at the peak of either depolarizing current wave or muscle stretching both preceded by a change in opposite direction (the spikes coinciding with the peak in muscle length PSA being actually elicited by muscle spindle action potentials triggered at the moment of the peak stretching velocity). Time coupling of action potentials occurred during both muscle stretching and transmembrane stimulation, being more tight in the latter case as well as when larger amplitudes of the stimuli were applied. It is supposed that discharges from muscle spindle primary endings phase-locked to small pseudo-random muscle length changes may, due to the time coupling of skeletomotor action potentials, provoke a synchronous firing of skeletomotor neurones, mostly of those belonging to slow motor units. Possible effects of such a firing pattern on the resulting muscle reflex contraction and the stretch reflex stability as well as a possibility of it being provoked by fusimotor discharges are discussed.     

Discharge characteristics of spindle receptors located in different parts of the cat soleus muscle

Discharge characteristics of 48 receptors of passive muscle spindles during gradual stretching of the soleus muscle were studied in anesthetized cats. The position of the receptor was determined by electrical stimulation of the muscle surface, by pressure on the muscle, by stretching of individual bundles of fibers, by intramuscular electrical stimulation, and also by coagulation of areas of muscle leading to disappearance of discharges of the monitored unit. Primary and secondary endings were found in which dynamic and static sensitivity could be distinguished and were independent of localization. Primary endings with identical functional properties were found in both the proximal and middle parts of the muscle. It is postulated that lower or higher dynamic and static sensitivity of receptors depends not on the location of muscle spindles in the soleus muscle, but on differences in the density of afferent terminal contacts with three types of intrafusal fibers.     

Changes in sensitivity of muscle spindles of fast and slowly contracting muscles after chronic de-efferentation

Activity of muscle spindles of fast (m. EDL) and slowly contracting (m. sol.) muscles was studied in cats 2–5 weeks after extradural division of the ventral roots of the spinal cord at the level of segments L₅–S₂. Spontaneous activity of muscle spindles recorded against the background of a relaxed muscle was unchanged after de-efferentation; spontaneous activity, however, evoked by stretching the muscles with a load of 100 g, was increased as a result of denervation in both primary and secondary endings. During additional stretching of the muscles with different speed and amplitude the sensory endings of the muscle spindles under conditions of chronic de-efferentation generated responses with higher than normal frequency in both the dynamic and the static phases of stretching. In this case a greater increase in frequency was observed in the slow than in the fast muscle.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 13, No. 2, pp. 204–209, March–April, 1981.     

Synchronization of motor units and its simulation in parallel feedback system

Comparison of the discharge patterns of soleus motor units with associated changes in force exerted by foot during quiet stance have already demonstrated the following facts. In the initial stage of standing, the motor units exhibited stationary and asynchronous discharges. The force showed a sporadic presence of the high frequency oscillation in the 8–10 Hz band. After five to ten minutes of standing, the firing rate of individual motor unit discharges increased to about 10 spikes/sec and discharges of each motor unit were synchronized and phase-locked to each of the accompanying force oscillation. During this transitional stage, the discharges of motor units were characterized by spike dropouts from an otherwise regular spike train. To simulate the changes in the discharge characteristics, we have proposed a parallel feedback model of the stretch reflex arc. This was made of multiple -motoneurons, motor units and muscle spindles. And motor units interact each other through group Ia afferent signals. As a result of simulation, motor units were found to exhibit stationary and asynchronous discharges when feedback gain was kept small. With an increase of feedback gain, the firing rate of individual motor units increased and finally the discharges of them were synchronized. During this transitional stage, the spike dropouts were observed in accordance with the experimental results. The neuronal mechanism of synchronization may partly be explained by the interactions of motor neurons through the above stated parallel feedback system.     

Effect of motor stimulation and stretching on afferent activity of the neuromuscular spindle isolated from the frog]

The arrangement of muscle spindles in m. ext. long. dig. IV has been examined by microdissection. It is confirmed that spindle systems generally appear to consist of individual receptors. Stimulation effects of fast motor fibres (conduction velocities greater than 12 m/sec) on the spindles of the same muscle were studied. Receptors were isolated with their nerves and the appropriate spinal roots, the latter ones were used for stimulating efferent fibres and recording sensory discharges. Single shocks to the ventral root filaments caused afferent responses ranging from a single action potential to a train of impulses. During repetitive stimulation (train of stimuli at frequency of 10 to 150/sec) a marked increase in afferent activity was found. Afferent activity could be driven by the frequency of stimuli ("driving") and the stimulus/action potentials ratio varied from 1:1 to 1:3 or more. The rate of sensory discharge depended on the frequency of stimuli: the maximum effect, was attained at 30 to 50 stimuli/sec and, in the most responsive receptors, up to 80 stimuli/sec. Slight increases of the initial lengths of the receptors caused facilitation of sensory responses to motor stimulation. Moreover, impairing effects, which appear during sustained or high-frequency stimulation, possibly related to fatigue in intrafusal neuromuscular transmission, could be relieved by increasing the initial length. The repetitive stimulation of fast fusimotor fibres increased both dynamic and static responses and also raised the afferent activity after a period of stretching, when usually a depression occurs; these effects varied according to the preparation, its initial tension and the frequency of stimulation. The main feature of the examined motor fibres, when stimulated, is the constant excitatory action on muscle spindle static response. Results are discussed. It is suggested that the different characteristics of intrafusal muscle fibres, the receptor initial tension and the frequency of motor units discharges, may together affect muscle spindles static or dynamic performance.     

Testing excitability in human firing motoneurons during the interspike interval

Monosynaptic testing of excitability in firing triceps surae muscle motoneurons activated during volitional contraction was performed using a technique for recording potentials from single motor units and by producing H-reflex. Motoneuronal excitability was assessed according to level of firing index. Motoneuronal firing index decreased during transition from a low background rhythmic firing rate of less than 6 spikes/sec to one of 6–8 spikes/sec. It hardly changed with a further rise in rate to 12 spikes/sec. The dependence between firing index and spike rate are put down to changes occurring in motoneuronal excitability during the interspike interval. Findings indicate that in the low frequency range of motoneuronal firing characteristic of natural muscle contraction, discharge rate may be considered one of the factors determining excitability in the motoneuron and hence its transmission qualities.Institute of Problems in Information Transmission, Academy of Sciences of the USSR. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 210–216, March–April, 1987.     

Comparative analysis of background activity of ventrolateral thalamic neurons in patients with parkinsonism and torsion muscle dystonia

In the course of neurosurgical interventions in 40 patients with parkinsonism and torsion muscle dystonia, the background activity (BA) was recorded from 124 neurons of the ventrolateral nucleus (VL) of the thalamus with the aid of microelectrodes during demarcation of boundaries between nuclear structures, and identification of zones within a nucleus. The following characteristic features of the BA in patients with parkinsonism were found: a relatively large proportion (71%) of cells with a burst pattern of activity; a great diversity of burst duration in the activity (short or prolonged bursts); a large proportion (67%) of cells with cyclic changes in the BA frequency; diverse patterns of cyclic modulation, where periods varied from fractions of a second (0.2–0.8 sec) to seconds (2–10 sec) or to tens of seconds (20–40 sec); cyclic successions of spike bursts with the rhythm typical of the peripheral tremor (3–7/sec) in a substantial proportion (40%) of the units.The background spike trains recorded in patients with torsion muscle dystonia consisted of separated spikes in the majority (69%) of units. Only short high-frequency discharges were found in the burst activity in this group; a cyclic BA pattern occurred much less frequently (in 23% of neurons); burst discharges at a 3–4/sec frequency were found only in 4% of the examined cells.The possible nature of motor disorders in patients with parkinsonism and torsion muscle dystonia and the mechanisms of the curing effects of cryodestruction of theVL of the thalamus in the treatment of the disease are discussed.Neirofiziologiya/Neurophysiology, Vol. 25, No. 4, pp. 246–253, July–August, 1993.     

Effect of thyrotoxicosis on the activity of proprioceptors of the skeletal muscles]

A study was made of the influence of prolonged administration of the thyroid hormones on the pulse activity of the cat soleus muscle spindles. In the animals with thyrotoxicosis the response of the primary endings of the spindles on the continuously acting and sudden rapid extension proved to be considerably enhanced both during the dynamic and the static phase. The frequency of discharges of the secondary endings showed no significant changes. It is supported that the changes in the activity of the primary endings observed were connected with the disturbances in the metabolism of the muscle and also with its atrophy.     

Investigation of the frequency distribution of natural activity in a mixed nerve trunk by the colliding impulse method

the theory of the colliding impulse method of determining the discharge frequency spectrum and the distribution of fibers by frequency in a mixed nerve trunk is examined. Using the double impulse method, a modification of the colliding impulse method, afferent A-fibers of the aortic nerve in the cat were investigated. Only 61% of these fibers were found to be activated in the initial phase of cardiac ejection when the arterial pressure was 120 mm Hg. Most of the active fibers functioned with a frequency of about 115 spikes/sec.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 6, No. 3, pp.318–327, May–June, 1974.     

Model study of muscle spindles subjected to static fusimotor activation

An earlier proposed mechanical model of the de-efferented muscle spindle endings has been developed further in order to study whether the functional effects of activating the static fusimotor fibres may be accounted for by relatively simple mechanical events in the intrafusal muscle fibres. Particular attention has been paid to important controversial problems related to the processes involved in the fusimotor activation of the mammalian muscle spindles. In order to develop an adequate model, preliminary simulation studies of the functional effects of various reasonable modifications of the original “de-efferented” model were first carried out by means of a convenient direct electronic analogue of the mechanical system. The following results apparently reflect pertinent details about the peripheral receptor mechanisms associated with the activation of the static fusimotor fibres:

1. The γ _s-activation is most adequately accounted for by mechanical events occurring in the nuclear chain fibres.
2. Any uniform chain fibre contraction fails to account for the effects of the γ _s-activation.
3. The simulation of a local chain fibre contraction could apparently account for all significant effects of the γ _s-activation.
4. A reasonable increase of the position sensitivity and the vibration sensitivity of both the primary and the secondary endings could be accounted for by increasing the stiffness of about 1/4–1/2 of the length of the simulated nuclear chain fibre by a factor of about 2–10.
5. The powerful modulation of the background discharge of the receptor endings that is associated with the γ _s-activation could be accounted for by an independent contraction force related to the intensity of the static fusimotor activity.
6. The model predicts a decrease of the rate sensitivity of the primary endings (group Ia) and a slight increase of the (moderate) rate sensitivity of the secondary endings.
7. By an adequate selection of the parameters, the simulated γ _s-activation showed a negligible influence on the absolute level of the dynamic phase of the ramp response of the simulated primary endings, in spite of a considerable increase of the steady state (background) discharge before and after the termination of a simulated stretch.

     

Correlation between discharges of human motor units during prolonged muscle contraction

Motor unit (MU) potentials were recorded from the human rectus femoris and biceps brachii muscles during prolonged isometric contraction. Interspike intervals and intervals between adjacent discharges of 2 MUs (cross-intervals of MU pairs) were measured. Synchronization was expressed by the following criteria: the cross-interval histogram; comparison of the number of coincidences between discharges of 2 MUs observed experimentally with the mean probable number of coincidences; the frequency of appearance of N successive coincidences of spikes from different MU pairs; comparison of the mean duration of interspike intervals preceding a synchronized discharge with the mean duration of the remaining interspike intervals for the same MU. For some MU pairs the number of coinciding spikes was greater than the expected number of random coincidences. Synchronized spikes could form a train of consecutive coincidences. The mean duration of interspike intervals preceding a synchronized discharge was somewhat less than the mean duration of the remaining interspike intervals for MUs forming a synchronously firing pair.Institute of Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 68–74, January–February, 1972.