Monosynaptic inhibitory postsynaptic potentials of cortical neurons

Monopolar intracortical stimulation of the auditory cortex was carried out in cats immobilized with D-tubocurarine. A macroelectrode (tip diameter 100 µ) or a microelectrode (tip diameter 10–15 µ) was used for stimulation. In both cases, besides excitatory responses, primary IPSPs with latent periods of 0.4–1.2 and 1.4–6.0 msec were recorded in cortical neurons close to the point of stimulation. The first group of IPSPs are considered to be generated in response to direct stimulation of bodies or axons of inhibitory cortical neurons, i.e., monosynaptically. The amplitude of these IPSPs varied in different neurons from 3 to 15 mV, and their duration from 4 to 150 msec. Additional later inhibitory responses were superposed on many of them. Of the IPSPs generated in auditory cortical neurons in response to stimulation of geniculocortical fibers 1.5% had a latency of 0.8–1.3 msec. They also are assumed to be monosynaptic. It is concluded that the duration of synaptic delay of IPSPs in cortical neurons and spinal motoneurons is the same, namely 0.3–0.4 msec. Axons of auditory cortical inhibitory neurons may be 1.5 mm long. The velocity of impulse conduction along these axons is 1.6–2.8 m/sec. The genesis of some special features of IPSPs of cortical neurons is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 5, pp. 458–467, September–October, 1975.     

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.     

Electrophysiological analysis of the functional organization of cortico-cerebellar connections

The effect of stimulation of cortical association (orbito-frontal, parietal) and projection (auditory, sensomotor) areas on the activity of Purkinje neurons of the cerebellar cortex was studied in adult cats anesthetized with pentobarbital, with or without chloralose. These responses were compared with those to peripheral stimuli. Definite similarity was found between the responses of Purkinje cells to different cortical (association and projection) stimuli as regards both the types of responses of the neurons and their ability to respond. No similarity was observed in the responses of Purkinje cells to peripheral (visual, auditory, electrodermal) stimulation. Whereas almost identical numbers of neurons (over 50%) were excited in response to the different forms of cortical stimulation, the ability of the neurons to respond to peripheral stimuli differed considerably: 44.6% of neurons responded to electrodermal stimulation, 34.2% to auditory, and 18.8% to visual.Medical Institute, Kemerovo. Translated from Neirofiziologiya, Vol. 8, No. 5, pp. 483–489, September–October, 1976.     

Responses of auditory cortex neurons in unanesthetized cats to best-frequency tones

The characteristics of extra- and intracellular responses of neurons in the AI region were studied in experiments with unanesthetized cats. It was established that auditory cortex neurons with similar best frequencies showed different forms of responses to tones of the corresponding frequency. About 40% of the auditory cortex neurons generated on responses to tone presentation. On — off and off responses were found in 27% of the neurons. Cortical neurons (27%) in which stimulation or inhibition of impulse discharge persisted throughout tone action were assigned to the tonic type group of cells. Approximately 6% of neurons in the AI region did not respond to a tone. During intracellular recording about 85% of the neurons responded to the turning on and/or off of a tone by generating an action potential followed by an IPSI. In 96% of the cortical neurons studied the IPSPs were a constant component of the intracellular responses to a tone. It is concluded that the inhibition of the impulse activity of the given neurons is of primarily a postsynaptic origin. Neurons showing one or another form of response differ from one another in the relative intensity and time characteristics of excitatory and inhibitory processes interacting on their postsynaptic membranes. In neurons of the phasic type inhibitory processes are dominant over excitatory, while excitatory processes are predominant in neurons of the tonic type.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 500–508, July–August, 1985.     

Responses of medial geniculate body neurons to auditory cortical stimulation

Extracellular and intracellular unit responses of thepars principalis of the medial geniculate body to stimulation of the first (AI), second (AII), and third (AIII) auditory cortical areas were studied in cats immobilized with D-tubocurarine. In response to auditory cortical stimulation both antidromic (45–50%) and orthodromic (50–55%) responses occurred in the geniculate neurons. The latent period of the antidromic responses was 0.3–2.5 msec and of the orthodromic 2.0–18.0 msec. Late responses had a latent period of 30–200 msec. Of all neurons responding antidromically to stimulation of AII, 63% responded antidromically to stimulation of AI also, confirming the hypothesis that many of the same neurons of the medial geniculate body have projections into both auditory areas. Orthodromic responses of geniculate neurons consisted either of 1 or 2 spikes or of volleys of 8–12 spikes with a frequency of 300–600/sec. It is suggested that the volleys of spikes were discharges of inhibitory neurons. Intracellular responses were recorded in the form of antidromic spikes, EPSPs, EPSP-spike, EPSP-spike-IPSP, EPSP-IPSP, and primary IPSP. Over 50% of primary IPSP had a latent period of 2.0–4.0 msec. It is suggested that they arose through the participation of inhibitory interneurons located in the medial geniculate body.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 1, pp. 5–12, January–February, 1976.     

Effect of the vermian cortex of the anterior lobe of the cerebellum on the background electrical activity of fastigial neurons

The effect of electrical stimulation of the vermian cortex of the anterior lobe of the cerebellum on the activity of neurons of different portions of the fastigial nucleus was studied in acute experiments on cats under light nembutal anesthesia. Inhibitory influences of the cortex (80.6% of the reacting neurons) were manifested in total blocking or decrease in the frequency of the background activity of the neurons and were characterized by a long aftereffect and "rebound." Stimulation of the cortex also had a blocking effect on the inhibitory interaction of adjacent nuclear elements. Facilitatory influences (14.5% of the neurons) were expressed either by an increase of spontaneous discharges of the neurons or by the appearance of activity in rhythm with the stimulation. The effectiveness of cortical stimulation depended on the localization of the stimulating electrodes. Zones of maximum density of projections to a given neuron of the nucleus and convergence and divergence of influences were found in the cerebellar cortex.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSR, Erevan. Translated from Neirofiziolgiya, Vol. 2, No. 3, pp. 260–268, May–June, 1970.     

Dynamics of functional reciprocal connections between cerebral motor cortex neurons under stimulation in cats

Activity was recorded from neurons belonging to the representation of the forelimb in the motor cortex (sulcus cruciatus, L 7–9 mm) using multiple multi-channel/barrel electrodes during acute experiments on cats. Cross-correlation analysis of impulse trains was adopted to investigate dynamics of interneuronal connections during passive flexion and electrical stimulation of the limb contralateral to the recording site. It was found that neither passive bending nor electrical stimulation of the limb leads to a significant increase in the total number of direct relationships between cortical neurons. At the same time, passive flexion does produce a considerable decrease in the number of instances of both inputs operating in neighboring neurons (50–100 µm apart) and an increase in cells located further (between 100 and 400 µm) apart. Some increase in the number of direct inhibitory interactions between neighboring neurons was observed during electrical stimulation.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Nentskii Institute of Experimental Biology, Warsaw, Poland. Center of Experimental and Clinical Medicine, Warsaw, Poland. Translated from Neirofiziologiya, Vol. 23, No. 1, pp. 73–80, January–February, 1991.     

Responses of rabbit visual cortical neurons to intracortical electrical stimulation

Responses of rabbit visual cortical neurons to single and repetitive intracortical electrical stimulation were investigated. The stimulating electrode was located 0.7–1.2 mm away from the recording electrode. Response thresholds to single stimulation were as a rule 150–180 µA, whereas to series of stimuli they were 30–60 µA. The latent period to the first spike averaged 5–15 msec but the probability of the initial discharge was very low, namely 3–6%. With an increase in current intensity the duration of the initial inhibitory pause was increased in half of the neurons responding to it, whereas in the rest it was unchanged. After presentation of series of stimuli spontaneous activity was enhanced for a short time (4–6 sec). In about half of the cells the same kinds of discharge dynamics were observed in response to repetitive stimulation (frequency 0.25 Hz) as in responses to light, but more neurons with sensitization of discharge and fewer "habituating" neurons took part in responses to electrical stimulation. It is postulated that stimulation of a given point of the visual cortex evokes excitation of a local neuron hypercolumn and inhibition of neighboring cell columns.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 15, No. 4, pp. 412–419, July–August, 1983.     

Interaction between neurons displaying tonic response to sound and other cells of the feline auditory cortex

Interaction between neurons with a tonic response pattern and either nearby or further removed (by about 400–500 µm) cortical neurons was investigated during acute experiments on 15 immobilized cats using cross-correlation analysis techniques. Synchronizing excitatory input common to the nerve cells was found in cross-correlation histograms (CCH) in 26 out of 36 test pairs of neurons (72%). Both positive and negative cross-correlation (five pairs in each case) were discovered, pointing to mono- or polysynaptic excitatory and inhibitory effects of the tonic neuron on spike activity in the other cell from CCH of 10 pairs of neurons. The functional diversity of neurons f distinguished by a tonic pattern of response to sound was deduced on the basis of findings from this research. The theory that some tonic type cells act as excitatory neurons and others fulfil the function of inhibitory interneurons is examined.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 5, pp. 613–620, September–October, 1989.     

Effect of intracortical chemical stimulation on spontaneous sensomotor cortical unit activity in cats

The effect of microelectrophoresis of glutamate on spontaneous activity of sensomotor cortical neurons located 80–250 µ from the point of application of glutamate was studied in cats anesthetized with pentobarbital. If glutamate was applied at distances of under 100 µ from the neurons the predominant response was one of excitation, evidently due to the direct action of the excitatory mediator. With more distant application inhibition of spontaneous activity predominated: at distances from 100 to 200 µ it was observed in 57%, and between 200 and 250 µ, in 70% of cases. Application of picrotoxin close to the neuron weakened inhibition induced by microelectrophoresis of glutamate through a neighboring microelectrode.A. A. Ukhtomskii Physiological Research Institute, A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 347–352, July–August, 1982.     

Tonotopic organization of the ventrorostral zone of the auditory cortex region AII in cats

The tonotopic organization of the ventrorostral (VR) zone of cortical auditory area AII was investigated in acute experiments on cats anesthetized with nembutal and unanesthetized immobilized animals. Response with the lowest threshold arose in 92% of test neurons to presentation of one or several sound frequencies. The majority (54%) were "tuned" to one characteristic frequency (CF), 38% to several frequencies, and 8% had no clear-cut CF. A connection was found between location of a unit within the VR zone and its CF. Neurons with the highest CF were located in the ventrocaudal AII. An increase was noted in numbers of neurons with the lowest CF with increasing distance (rost-rally) from the VR location zone of neurons tuned to a high frequency. Going by response to acoustic stimuli of frequencies ranging between 1 and 24 kHz, length of the VR projection zone of the AII was found to measure 1.8–2.0 mm. Location of the test zone in relation to auditory cortex sulci varied substantially from one animal to the next.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 2, pp. 178–184, March–April, 1990.     

Neuronal responses of a chronically isolated slab of auditory cortex to intracortical stimulation during paroxysmal electrical activity in cats

Responses of 155 neurons 3 weeks after neuronal isolation of a slab of auditory cortex (area AI) to single intracortical stimulating pulses at the level of layer IV were studied in unanesthetized, curarized cats during paroxysmal electrical activity evoked by series of high-frequency (10–20 Hz) electrical stimulation by a current 2–5 times above threshold for the direct cortical response. In response to such stimulation a discharge of paroxysmal electrical activity, lasting from a few seconds to tens of seconds, appeared in the slab. As a rule it consisted of two phases — tonic and clonic. This indicates that cortical neurons can form both phases of paroxysmal cortical activity. Depending on behavior of the neurons during paroxysmal electrical activity and preservation of their ability to respond to intracortical stimulation at this time, all cells tested in the isolated slab were divided into four groups. Their distribution layer by layer and by duration of latent periods was studied. Two-thirds of the neurons tested were shown to generate spike activity during paroxysmal discharges whereas the rest exhibited no such activity. A special role of neurons in layer II in generation of paroxysmal activity in the isolated slab was noted. The view is expressed that at each moment functional neuronal circuits, independent of each other, exist in the slab and also, evidently in the intact cortex, which can interact with one another when conditions change.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 3–11, January–February, 1984.     

Organization of the Connections between the Parietal Associative Zone and the Primary Sensory Zones in the Cat Neocortex

In acute experiments on cats, we studied the impulse activity of 262 neurons of the parietal associative zone (PAZ, field 5). Among them, 129 cells [100 silent units and 29 units generating background activity (BA)] were identified as output neurons, while 133 cells with the BA were interneurons of the intrinsic cortical neuronal circuits. Electrical stimulation of the primary visual, auditory, or somatosensory cortices evoked no impulse responses in silent output PAZ neurons, while output neurons with the BA and interneurons (more than 65 and 80% of the cell units, respectively) generated clear responses (more frequently, phasic). Stimulation of the auditory and visual cortices exerted mostly inhibitory effects, while stimulation of the somatosensory cortex provided mostly excitatory influences. The ratios of neurons generating primary excitatory and inhibitory responses to stimulation of the visual, auditory, and somatic cortices were 0.3:1, 0.6:1, and 3.2:1, respectively. More than 95% of the field-5 neurons were influenced from the primary sensory zones via di- and/or polysynaptic pathways. Monosynaptic excitatory inputs from the visual cortex were identified for 3.8% of interneurons and 6.9% of output PAZ neurons; for the auditory cortical inputs, the respective figures were 1.7 and 3.5%. Monosynaptic connections with the somatic cortex were found only for 4% of the interneurons under study. It has been concluded that interaction of heteromodal signals coming to the PAZ via the corticopetal and associative inputs occurs on neurons of all the cortical layers.     

Single unit activity related to acoustic stimulus meaning in the cat auditory cortex during instrumental food reflex

Responses of 93 neurons to isolated presentation of a single click and a series of 10 clicks with following frequency of 1000 Hz and responses of 66 neurons after the click had become a positive conditioned stimulus, and a series of 10 clicks had become a differential, negative stimulus, were investigated in chronic experiments on cats. Formation and realization of differential inhibition of an instrumental food reflex was shown not to lead to strengthening of inhibition in the auditory cortex, and the process of differential inhibition itself within the primary auditory cortex is not essentially an inhibitory process. Identical changes were found in responses of auditory cortical neurons to both positive and negative conditioned stimuli after training, evidence that neurons of the primary auditory cortex play a similar role in realization of the instrumental reflex and in its differential inhibition. It is suggested that the presence of groups of neurons responding by excitation or inhibition only to presentation of a stimulus with definite informative value is of great importance for differentiation.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukranian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 2, pp. 212–221, March–April, 1985.     

Postsynaptic inhibition in the general cortex of the turtle forebrain

Postsynaptic inhibition in the general cortex of the turtle forebrain was investigated by recording unit activity intracellularly. Depending on the type of IPSPs recorded in response to electrical stimulation of the contralateral optic nerve and cortical surface the neurons were subdivided into three groups: 1) with long direct IPSPs, 2) with long and short direct, and also recurrent IPSPs, 3) with short direct and recurrent IPSPs. It is concluded that inhibitory pathways of the short direct and recurrent IPSPs have a common final component, a stellate interneuron. Compared with the recurrent collaterals of the principal neurons, the direct afferents make contact with more distal portions of the dendrites of this cell. Synapses formed on dendrites of the principal neurons by axons of the stellate cells are nearer to the soma than synapses responsible for generation of the long direct IPSP.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 5, No. 4, pp. 375–383, July–August, 1973.     

Evoked impulse activity in neuronally isolated cortex

The reactions of neurons of the isolated cortex of one hemisphere to direct cortical stimulation were investigated in cats under Nembutal anesthesia. Isolation of the cortex was carried out by Khananashvili's method [10]. It is shown that phasic reactions develop in the isolated cortex in response to such stimulation: initial discharge, initial pause, first after-discharge, first after-pause, late after-discharges and pauses, as well as reactions of presumably inhibitory neurons. A majority of the cells (85%) which manifest background activity respond to direct electrical stimulation, and the frequency of the late after-reactions is twice as great as in the intact cortex. It is concluded that cortical elements of the isolated cortex retain their principal neurophysiological properties.Institute of Experimental Medicine, Academy of Medical Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 3, No. 3, pp. 236–244, May–June, 1971.     

Unit responses in the cat auditory cortex to medial geniculate body stimulation

Responses of 98 auditory cortical neurons to electrical stimulation of the medial geniculate body (MGB) were recorded (45 extracellulary, 53 intracellularly) in experiments on cats immobilized with tubocurarine. Responses of the same neurons to clicks were recorded for comparison. Of the total number of neurons, 75 (76%) responded both to MGB stimulation and to clicks, and 23 (24%) to MGB stimulation only. The latent period of extracellularly recorded action potentials of auditory cortical neurons in response to clicks varied from 7 to 28 msec (late responses were disregarded), and that to MGB stimulation varied from 1.5 to 12.5 msec. For EPSPs these values were 8–13 and 1–4 msec respectively. The latent period of IPSPs arising in response to MGB stimulation varied from 2.2 to 6.5 msec; for 34% of neurons it did not exceed 3 msec. The difference between the latent periods of responses to clicks and to MGB stimulation varied for different neurons from 6 to 21 msec. Responses of 11% of neurons to MGB stimulation, recorded intracellularly, consisted of sub-threshold EPSPs, while responses of 23% of neurons began with an EPSP which was either followed by an action potential and subsequent IPSP or was at once cut off by an IPSP; 66% of neurons responded with primary IPSPs. Neurons responding to MGB stimulation by primary IPSPs are distributed irregularly in the depth of the cortex: there are very few in layers III and IV and many more at a depth of 1.6–2 mm. Conversely, excited neurons are predominant in layer III and IV, and they are few in number at a depth of 1.6–2 mm. It is concluded that the afferent volley reaching the auditory cortex induces excitation of some neurons therein and, at the same time, by the principle of reciprocity, induces inhibition of others. This afferent inhibition takes place with the participation of inhibitory interneurons, and in some cells the inhibition is recurrent. The existence of reciprocal relationships between neurons in different layers of the auditory cortex is postulated.A. A. Bogomolets' Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 4, No. 1, pp. 23–31, January–February, 1972.     

Electrophysiological investigation of conduction of afferent impulses through the medial geniculate body

Extra- and intracellular reactions of 280 neurons of the pars principalis of the medial geniculate body (MGB) and of 408 auditory cortical neurons in area AI to stimulation of the inferior brachium of the midbrain and geniculocortical fibers were studied in cats immobilized with D-tubocurarine. Single electrical stimulation of the inferior brachium was shown to evoke a long and complex neuronal response in MGB in the form of excitation of some and inhibition of other neurons. The initial component of this response lasted 13 msec. Excitation of 72% of neurons participating in the response took place during the first 3 msec after the beginning of stimulation. In the same period 84% of IPSP arose. The inferior brachium was shown to contain a certain number of descending fibers. Some of them are axons of MGB neurons. Many fibers of the inferior brachium reach the auditory cortex without synaptic relay in MGB. Of all cells of MGB excited by stimulation of the inferior brachium monosynaptically, 76% are thalamocortical relay neurons; the rest are interneurons. Of the relay neurons of MGB 90% are excited monosynaptically, the rest by impulses passing through two or three synaptic relays in MGB. During stimulation of the inferior brachium, responses consisting of EPSP-IPSP and primary IPSP are recorded in many neurons of MGB. About 20% of primary IPSP arise monosynaptically, evidently in response to stimulation of inhibitory fibers of the inferior brachium. Most IPSP arise disynaptically, with the participation of an inhibitory interneuron located at the entrance to MGB. Inhibition observed in this case is direct afferent in nature.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 11, No. 6, pp. 515–523, November–December, 1979.     

Unit responses in the cat auditory cortex to electrical stimulation of nerve fibers innervating receptor cells in different parts of the organ of corti

Responses of 200 primary auditory cortical neurons to electrical stimulation of nerve fibers in different receptor zones of the cochlea were studied in cats anesthetized with pentobarbital. Under the influence of paired stimulation, after the response to the conditioning stimulus a state of prolonged (from 4 to 200 msec) refractiveness to the second stimulus developed in all the neurons tested. This long-lasting inhibition of unit activity was due to inhibition developing in the thalamus and the auditory cortex itself. The intensity and duration of excitation and inhibition in the cortical projection focus were maximal when the center of the receptive field was stimulated and decreased when the stimulus shifted from the center to the periphery. The region of the receptor surface of the cochlea to stimulation of which the auditory cortical neurons respond by an action potential is much narrower than the region whose electrical stimulation depresses the discharge of these neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 418–425, July–August, 1982.     

Mechanisms of the effect of diazepam on paroxysmal electrical activity of an isolated cortical slab in cats

The effect of diazepam on paroxysmal global electrical activity of a neuronally isolated slab of auditory cortex and on inhibitory responses of its neurons due to intracortical electrical stimulation was investigated in cats. Diazepam (2 mg/kg, intravenously) caused inhibition of paroxysmal electrical activity and increased the number of inhibited neurons in both the acutely isolated slab and three weeks after isolation, compared with the intact cortex. However, the number of disynaptic responses was reduced under these circumstances, especially in the long-isolated slab. It is postulated that diazepam exerts its action through activation of GABA-ergic inhibitory neurons, by synchronizing inhibition and increasing the duration of the IPSPs. The action of diazepam is manifested first, probably, in the initial links of cortical neuron chains.I. I. Mechnikov Odessa State University. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 3–10, January–February, 1985.