Synaptic potentials of digastric-muscle motoneurons

We studied the antidromic and synaptic potentials evoked from 32 digastric-muscle motoneurons by stimulation of the motor nerve to this muscle, different branches of the trigeminal nerve, and the mesencephalic trigeminal nucleus. Antidromic potentials appeared after 1.1 msec and lasted about 2.0 msec. Stimulation of the infraorbital, lingual, and inferior alveolar nerves led to development of excitatory postsynaptic potentials (EPSP) and action potentials in the motoneurons. The antidromically and synaptically evoked action potentials of the digastric-nerve motoneurons were characterized by weak after-effects. We were able to record EPSP and action potentials in two of the motoneurons investigated in response to stimulation of the mesencephalic trigeminal nucleus, the latent period being 1.3 msec. This indicates the existence of a polysynaptic connection between the mesencephalic-nucleus neurons and the digastric-muscle motoneurons. Eight digastric-muscle motoneurons exhibited inhibitory postsynaptic potentials (IPSP), which were evoked by activation of the afferent fibers of the antagonistic muscle (m. masseter). The data obtained indicate the presence of reciprocal relationships between the motoneurons of the antagonistic muscles that participate in the act of mastication.A. A. Bogomol'ts Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 1, pp. 52–57, January–February, 1971.     

Role of different types of fibers of the infraorbital nerve in synaptic activation of masseter and digastric motoneurons

Postsynaptic potentials of motoneurons of the masseter and digastric muscles evoked by stimulation of the infraorbital nerve with a strength of between 1 and 10 thresholds were investigated in cats anesthetized with a mixture of chloralose and pentobarbital. Depending on their ability to be activated by low-threshold afferents of this nerve, motoneurons of the masseter were divided into two groups. Stimuli with a strength of 1.2–2.5 times above threshold for the most excitable fibers of the infraorbital nerve evoked short-latency EPSPs in the motoneurons of the first group; a further increase in stimulus strength (3–9 thresholds) led to the appearance of IPSPs with latent periods of 2.8–3.5 msec. Motoneurons of the second group responded to stimulation of the infraorbital nerve with a strength of 3–9 thresholds by IPSPs whose latent periods varied from 6 to 8 msec. Stimuli below 3 thresholds in strength evoked no responses in these motoneurons. Stimulation of the infraorbital nerve with pulses of between 1 and 2 thresholds in strength evoked EPSPs in digastric motoneurons, but an increase in the strength of stimulation led to action potential generation. The presence of many excitatory and inhibitory inputs formed by afferent fibers of different types evidently provides a basis for functional diversity of jaw-opening and jaw-closing reflexes.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 6, pp. 596–603, November–December, 1980.     

Membrane electrical properties of motoneurons innervating the masseter muscles in rats

Membrane potentials and action potentials evoked by antidromic and direct stimulation were investigated in motoneurons of the trigeminal nucleus in rats innervating the masseter muscle. This motor nucleus was shown to contain cell populations with high and low membrane potentials. The responses of cells of the first group had shorter latent periods of their antidromic action potentials, a longer spike duration, and a lower amplitude and shorter duration of after-hyperpolarization than responses of cells of the second group, and the input resistance of their membrane also is lower. The bimodal character of distribution of electrophysiological parameters of motoneurons in the trigeminal nucleus indicates that "fast" and "slow" fibers of the masseter muscles may be innervated by different types of nerve cells.N. A. Semashko Moscow Medical Stomatological Institute. Translated from Neirofiziologiya, Vol. 13, No. 3, pp. 270–274, May–June, 1981.     

Corticofugal influences on the motoneurons of the trigeminal nucleus

We studied the postsynaptic potentials evoked from 76 trigeminal motoneurons by stimulation of the motor (MI) and somatosensory (SI) cortex in the ipsilateral and contralateral hemispheres of the cat. Stimulation of these cortical regions evoked primarily inhibitory postsynaptic potentials (PSP) in the motoneuron of the masseter muscle, but we also observed excitatory PSP and mixed reactions of the EPSP/IPSP type. The average IPSP latent period for the motoneurons of the masseter on stimulation of the ipsilateral cortex was 6.1±0.3 msec, while that on stimulation of the contralateral cortex was 5.2±0.4 msec; the corresponding figures for the EPSP were 7.6±0.5 and 4.5±0.3 msec respectively. Corticofugal impulses evoked only EPSP and action potentials in the motoneurons of the digastric muscle (m. digastricus). The latent period of the EPSP was 7.6 msec when evoked by afferent impulses from the ipsilateral cortex and 5.4 msec when evoked by pulses from the contralateral cortex. The duration of the PSP ranged from 25 to 30 msec. Postsynaptic potentials developed in the motoneurons studied when the cortex was stimulated with a single stimulus. An increase in the number of stimuli in the series led to a rise in the PSP amplitude and a reduction in the latent periods. When the cortex was stimulated with a series of pulses (lasting 1.0 msec), the IPSP were prolonged by appearance of a late slow component. We have hypothesized that activation of the trigeminal motoneurons by corticofugal impulsation is effected through a polysynaptic pathway; each functional group of motoneurons is activated in the same manner by the ipsilateral and contralateral cortex. The excitation of the digastric motoneurons and inhibition of the masseter motoneurons indicates reciprocal cortical control of their activity.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 3, No. 5, pp. 512–519, September–October, 1971.     

EPSPs of masseter motoneurons evoked by stimulation of low-threshold infraorbital nerve afferents in cat

Stimulation of the infraorbital nerve at strengths 1.4–2.5 times higer than the threshold of excitation of A fibers in cats anesthetized with chloralose and pentobarbital evoked EPSPs with an amplitude up to 3.0 mV and a duration of 9–15 msec in 69% of masseter motoneurons after 1.5–3.0 msec. These EPSPs were complex and formed by summation of simpler short-latency and long-latency EPSPs. The short-latency EPSPs appeared in response to infraorbital nerve stimulation at 1.1–1.5 thresholds and had a slow rate of rise (2.5–4.5 msec, mean 3.7±0.4 msec), low amplitude (under 2.0 mV), and short duration (5–6 msec). Their latent period varied from 1.5 to 3.0 msec (mean 2.1±0.2 msec). The shortness of the latent period and its constancy during stimulation of the nerve at increasing strength, and also the character of development of facilitation and inhibition of the EPSP during high-frequency stimulation suggests that these EPSPs are monosynaptic. The slow rate of rise suggested that these EPSPs arise on distal dendrites of the motoneurons. Long-latency EPSPs appeared 7–9 msec after stimulation of the infraorbital nerve at 1.1–1.5 thresholds. Their amplitude reached 1.5–2.0 mV and their duration 7–9 msec. The long duration of the latent period combined with low ability to reproduce high-frequency stimulation (up to 30/sec) points to the polysynaptic origin of these EPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 9, No. 6, pp. 583–591, November–December, 1977.     

Rubrofugal activation of motoneurons of the cat accessory nucleus

Postsynaptic potentials evoked in accessory nerve motoneurons by stimulation of the ipsilateral and contralateral red nuclei were investigated in acute experiments on cats anesthetized with chloralose and pentobarbital. Polysynaptic EPSPs with latent periods of 5.2 to 16 (mean 9.1 ± 0.7) msec and from 5.5 to 18 (mean 10.3 ± 0.9) msec, respectively, appeared in motoneurons of the accessory nerve in response to stimulation of the contralateral and ipsilateral red nuclei. A minimum of two or three stimuli was necessary to produce EPSPs in these motoneurons. In response to single stimulation of the contralateral and ipsilateral red nuclei EPSPs appeared in four motoneurons of the trapezius muscle with latent periods of 2.5 to 5.0 and 3.0 to 5.2 msec, respectively. An increase in the number of stimuli led to action potential generation by motoneurons. The functional role of such activation is discussed.A. A. Bogomolets Institue of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 5, pp. 532–536, September–October, 1982.     

Monosynaptic excitation of motoneurons of the accessory nerve nucleus by reticulofugal impulses

Acute experiments on cats anesthetized with chloralose and pentobarbital showed that excitation of fast-conducting (130 m/sec) reticulospinal fibers, arising during stimulation of the ipsilateral medullary reticular gigantocellular nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nerve nucleus. The EPSPs had latent periods of between 0.6 and 1.0 msec (mean 0.7 msec), they reached their maximal amplitude (4.0 mV) after 2.0–2.5 msec, and lasted about 10 msec. The EPSPs underwent only weak potentiation through the different types of stimulation of the gigantocellular nucleus and were not transformed into action potentials.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 12, No. 1, pp. 62–66, January–February, 1980.     

Postsynaptic potentials of facial motoneurons evoked by afferent and corticofugal impulses

Postsynaptic potentials of motoneurons in the facial nerve nucleus, evoked by stimulation of the cranial nerves (trigeminal, hypoglossal, facial) and of the sensomotor cortex were investigated in cats anesthetized with chloralose and pentobarbital. Two functionally opposite groups of motoneurons were found to exist in the facial nucleus. Stimulation of the afferent nerves and cortex evoked the appearance of EPSPs in the first of these groups and IPSPs in the second. The latency and duration of the PSPs indicate that afferent and corticofugal impulses reach the facial motoneurons along polysynaptic pathways. Interneurons on which wide convergence of influences travelling along afferent fibers and of the cortex, were found in the region of the facial nucleus. The possible neuronal pathways concerned with the transmission of afferent and corticofugal impulses to the facial motoneurons are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol.4, No.4, pp. 391–400, July–August, 1972.     

Neuromechanisms of reciprocal interrelation between jaw-opening and jaw-closing muscles in the cat (author's transl)]

Neural controlling mechanisms between the digastric (jaw-opening) and masseter (jaw-closing) muscles were studied in the cat. High threshold afferent impulses from the anterior belly of the digastric muscle to masseteric montoneurons in the trigeminal motor nucleus induced an EPSP-IPSP sequence of potentials with long latency, and high threshold afferent impulses from the masseter muscle also exerted a similar effect on digastric motoneurons in the same nucleus innervating the anterior belly of the digastric muscle. These results suggest that reciprocal inhibition via Ia interneurons as observed between the flexor and extensor muscles in the spinal cord does not exist between the digastric and masseter muscles in the cat. However, the respective motoneurons innervating the masseter and digastric muscles receive inputs of early excitation-late inhibition via high threshold afferent nerve fibers from each antagonistic muscle. As such, since EPSPs preceding IPSPs are recognized, these high threshold afferent impulses may exert not only a reciprocal inhibitory effect, but also a synchronous excitatory or inhibitory effect on the antagonistic motoneurons.     

Excitation of accessory motoneurons by stimulation of Deiters' nucleus in the cat

Acute experiments on cats under chloralose-pentobarbital anesthesia showed that application of single stimuli to Deiters' nucleus evoked monosynaptic EPSPs in motoneurons of the accessory nucleus. Latent periods of EPSPs ranged from 1.3 to 2.3 msec (mean 1.8±0.3 msec), their rise time was 0.5–1.0 msec, and their duration 7–10 msec. During repetitive stimulation the EPSPs were weakly potentiated, but with an increase in the strength of stimulation applied to Deiters' nucleus they readily changed into action potentials. In some motoneurons polysynaptic EPSPs with latent periods of the order of 6.0 msec appeared on the descending phase of these EPSPs.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 13, No. 5, pp. 515–519, September–October, 1981.     

Postsynaptic potentials of neck muscle motoneurons evoked by horizontal semicircular canal stimulation in cats

Postsynaptic potentials evoked by stimulation of ipsilateral and contralateral horizontal semicircular canals in motoneurons of muscles tilting and turning the head were investigated in acute experiments on cats anesthetized with chloralose and pentobarbital. Stimulation of the ipsilateral canal evoked EPSPs with latent periods varying from 1.8 to 10.0 msec in 25 of these motoneurons and IPSPs with latent periods varying from 1.9 to 3.9 msec in 10 of them. Calculation of the impulse conduction time from the ipsilateral semicircular canal through Deiters' nucleus to the cervical motoneurons indicates that EPSPs with latent periods of under 3.8 msec may be regarded as disynaptic, and those with latent periods of over 3.8 msec as polysynaptic. Stimulation of the contralateral canal evoked EPSPs with latent periods varying from 1.8 to 6.0 msec in 19 motoneurons and IPSPs with latent periods varying from 3.2 to 3.9 msec in two cells. The possible pathways of transmission of these influences and their functional role are discussed.     

Synaptic processes in facial motoneurons evoked by stimulation of the caudal trigeminal nucleus

Field potentials and postsynaptic potentials of facial motoneurons evoked by stimulation of the caudal trigeminal nucleus were investigated in acute experiments on cats by extra- and intra-cellular recording. Pre- and postsynaptic components of field potentials were found. Four types of motoneuron response were distinguished: EPSP with generation of single action potentials; a gradual shift of depolarization inducing grouped action potentials; a rhythmic discharge of action potentials arising at a low level of depolarization; and EPSPs or EPSP-IPSP sequences. The monosynaptic and (chiefly) polysynaptic nature of these responses was demonstrated. The possible mechanism of afferent control over facial motoneurons are discussed.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziologiya, Vol. 12, No. 3, pp. 272–282, May–June, 1980.     

Functional properties of interneurons of the masticatory reflex

Interneurons of the supratrigeminal nucleus, transmitting effects from the sensory and motor branches of the trigeminal nerve to motoneurons of the muscles of mastication were investigated. Two groups of interneurons with different functional connections were found. The first group (A) contains neurons excited during stimulation of the sensory branches and the motor nerve to the digastric muscle (A₁), neurons excited during stimulation of sensory branches and high-threshold afferents of the motor nerve to the masseter muscle (A₂), and neurons excited only by low-threshold afferents of the motor nerve to the masseter muscle (A₃). Neurons of the second group (B) were activated only by sensory fibers of the trigeminal nerve. It is postulated that interneurons of group A transmit inhibitory effects to motoneurons of antagonist muscles of the lower jaw. Group B interneurons participate in the transmission of excitatory influences to motoneurons of the digastric muscle.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 2, pp. 150–157, March–April, 1972.     

Detection and analysis of recurrent inhibition of firing motoneurons in man

During regular firing of "small" motor units, activated during weak voluntary contraction of the human soleus muscle, thick efferent fibers of n. tibialis were stimulated (a small M response was evoked, in which the small units did not participate). Peristimulus histograms of potentials of single motor units were constructed and the effect of stimulation on interspike interval duration was analyzed. The firing rate of the motor units was 4.5–7.6 spikes/sec. Stimulation of the nerve led to a sharp decrease in probability of their discharge or even complete temporary cessation of firing, i.e., it had a well marked inhibitory effect (lasting 10–20 msec). The latent period of inhibition (35–40 msec) was only a little longer than the latent period of the monosynaptic reflex of the soleus muscle. The effect of an inhibitory volley on duration of the interspike interval of the motor units depended on the time when the volley arrived during the interval. Lengthening of the interval was observed only if the inhibitory volley arrived in the second half or at the end of the interval. It is concluded that inhibition of firing of small motor units is due to Renshaw cells, activated on stimulation of axons of large motoneurons. The efficiency of a short (compared with the duration of the interspike interval) inhibitory volley reaching a motoneuron firing at low frequency characteristic of its adequate activation, is discussed.Institute for Problems of Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 88–96, January–February, 1984.     

Role of propriospinal neurons in inhibition of the afferent flow

The effects of stimulation of the dorsal funiculus on dorsal surface potentials (DSPs) of the spinal cord evoked by stimulation of a peripheral nerve and on antidromic action potentials (AAPs) evoked by stimulation of terminal branches of primary afferent fibers and recorded from the afferent nerve or dorsal root, were investigated in acute experiments on spinal cats and on cats anesthetized with pentobarbital and chloralose. Stimulation of the dorsal funiculus led to biphasic inhibition of the N₁-component of the DSP with maxima at the 15th–30th and 60th–80th milliseconds between the conditioning and testing stimuli. Maximal reinforcement of the AAP was found with these intervals. Bilateral division of the dorsal funiculi between the point of application of the conditioning stimuli and the point of recording the DSP abolished the first wave of inhibition of the DSP and the reinforcement of the AAP. After total transection of the cord above the site of conditioning stimulation the picture was unchanged. It is concluded that the initial changes in DSP and AAP are due to activation of the presynaptic inhibition mechanism by antidromic impulses traveling along nerve fibers running in the dorsal funiculus. Repeated inhibition of the DSP, like reinforcement of the AAP, can possibly be attributed to activation of similar inhibitory mechanisms through the propriospinal neurons of the spinal cord.Dnepropetrovsk State University. Translated from Neirofiziologiya, Vol. 5, No. 4, pp. 401–405, July–August, 1973.     

Postsynaptic potentials of motoneurons evoked by rubrofugal impulsation in the hypoglossal nucleus of the cat

The effect of stimulation of the ipsilateral and contralateral red nuclei on motoneurons of the hypoglossal nucleus was studied in cats anesthetized with chloralose and pentobarbital. In 35 (69%) of the 51 motoneurons tested, PSPs were generated in response to stimulation of the red nuclei by series of 3 to 5 stimuli of threshold strength and with a frequency of 500–600/sec. Of this number, 33 motoneurons responded to stimulation by EPSPs, whose latent periods varied from 3.5 to 14.0 msec (mean value for the ipsilateral red nucleus 5.7±0.75, for the contralateral nucleus 6.8±0.8 msec), whereas two motoneurons responded (after 6.2 msec) by IPSPs. Of the 35 motoneurons responding to stimulation of the red nuclei, stimulation of the lingual nerve evoked EPSPs in 31 and IPSPs in 4 (two of them were inhibited by rubrofugal impulses). IPSPs were generated as a result of stimulation of the lingual nerve in 16 motoneurons which did not respond to rubrofugal impulses.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 62–66, January–February, 1978.     

Postsynaptic effects evoked by activation of long propriospinal pathways in cervical spinal neurons in cats

Effects induced in motoneurons and interneurons of the cervical enlargements of the cat spinal cord by stimulation of the lateral and ventral funiculi at the lower thoracic level were studied under conditions producing degeneration of fibers of descending brain systems. Stimulation of this sort evoked PSPs (mainly of mixed character) in 57 of 90 motoneurons tested. In nine motoneurons the primary response consisted of monosynaptic EPSPs evoked by activity of fibers of the lateral funiculus, and in the rest it consisted of polysyanptic (at least disynaptic) EPSPs and IPSPs. Polysynaptic effects arising in the neuron in response to stimulation of the lateral and ventral funiculi usually differed only quantitatively. The intensity of excitatory synaptic action on motoneurons of the proximal muscle (especially thoracid) was much greater than that on motoneurons of distal muscles. Nearly all motoneurons with no synaptic action belonged to the latter group. Stimulation of the lateral and ventral funculi facilitated synaptic action induced in motoneurons by stimulation of high-threshold segmental afferents and led to excitation of interneurons located in the vectral quadrant, and had no effect on interneurons in the dorsal regions of gray matter. These effects are regarded mainly as the result of excitation of long ascending propriospinal pathways in the cervical parts of the cord; it is also postulated that some of them are evoked by the arrival of activity along collaterals of descending propiospinal pathways to the neurons in this region.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 4, pp. 339–347, July–August, 1979.     

Cortico-pyramidal and cortico-extrapyramidal synaptic influences on lumbar motoneurons in monkeys

Postsynaptic potentials evoked by stimulation of the motor cortex or pyramids before and after acute pyramidotomy were investigated in the lumbar motoneurons of monkeys. In response to activation of fibers of the pyramidal tract monosynaptic EPSPs predominated in motoneurons innervating the distal muscles of the hind limbs. Monosynaptic EPSPs in the motoneurons of the distal muscles had a significantly higher amplitude and could be evoked by weaker stimuli than EPSPs in the motoneurons of the proximal muscles. Cortico-motoneuronal EPSPs in the motoneurons of the distal muscles had a less marked frequency potentiation than EPSPs with monosynaptic segmental delay in the motoneurons of the proximal muscles. Cortico-extrapyramidal synaptic responses appeared in the pyramidotomized monkeys during intensive repetitive stimulation of the motor cortex in motoneurons of both distal and proximal muscles. These effects, transmitted by descending projections of the brain stem, may be responsible for the partial preservation of cortical motor control after pyramidotomy.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 4, No. 6, pp. 587–596, November–December, 1972.     

Synaptic mechanisms of central control of motoneuronal activity in the facial nerve nucleus

The responses of motoneurons of the facial nerve nucleus (FNN), evoked by stimulations of the oculomotor nerve nucleus, Edinger-Westphal's nucleus,substantia nigra, and entopeduncular nucleus, were studied in acute experiments on anesthetized and immobilized cats. The FNN motoneurons were identified by their antidromic activation after stimulation of various branches of the facial nerve. Stimulation of the oculomotor nerve nucleus, Edinger-Westphal's nucleus, and ipsi- or contralateral parts of thesubstantia nigra evoked mono- and polysynaptic EPSP in the FNN motoneurons, while stimulation of the entopeduncular nucleus elicited only polysynaptic EPSP. The influences from the above structures were shown to converge on the same FNN motoneurons. The findings are discussed considering morphological peculiarities of the afferent inputs to the FNN.Neirofiziologiya/Neurophysiology, Vol. 27, No. 2, pp. 116–125, March–April, 1995.     

Neuronal and synaptic mechanisms of cortical inhibition

The latent periods, amplitude, and duration of IPSPs arising in neurons in different parts of the cat cortex in response to afferent stimuli, stimulation of thalamocortical fibers, and intracortical microstimulation are described. The duration of IPSPs evoked in cortical neurons in response to single afferent stimuli varied from 20 to 250 msec (most common frequency 30–60 msec). During intracortical microstimulation of the auditory cortex, IPSPs with a duration of 5–10 msec also appeared. Barbiturates and chloralose increased the duration of the IPSPs to 300–500 msec. The latent period of 73% of IPSPs arising in auditory cortical neurons in response to stimulation of thalamocortical fibers was 1.2 msec longer than the latent period of monosynaptic EPSPs evoked in the same way. It is concluded from these data that inhibition arising in most neurons of cortical projection areas as a result of the arrival of corresponding afferent impulsation is direct afferent inhibition involving the participation of cortical inhibitory interneurons. A mechanism of recurrent inhibition takes part in the development of inhibition in a certain proportion of neurons. IPSPs arise monosynaptically in 2% of cells. A study of responses of cortical neurons to intracortical microstimulation showed that synaptic delay of IPSPs in these cells is 0.3–0.4 msec. The length of axons of inhibitory neurons in layer IV of the auditory cortex reaches 1.5 mm. The velocity of spread of excitation along these axons is 1.6–2.8 msec (mean 2.2 msec).A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 16, No. 3, pp. 394–403, May–June, 1984.