The spike reactions of the motor cortex neurons of old rabbits to specific stimuli

Spike reactions of motor cortex neurons to tactile and electrocutaneous stimulation of a forelimb were studied in aged (6-7-year old) rabbits. As compared with young adult animals, the neuronal reactions to afferent stimuli were rarely recorded in the motor cortex of aged rabbits (66.7 and 50%, respectively). The activation manifested in increasing firing rate over its spontaneous level was less intensive than in young animals. The neuronal reactions of aged animals were characterized by the slower activation with longer latencies and slower development of spike responses. The parameters of slow activation could be partly corrected by the iontophoretic application of acetylcholine to the soma region. Neuronal inhibition recorded in the motor cortex of aged rabbits was not markedly changed compared to inhibition reactions in young animals. It is suggested that impairment of the functional state of dendrites in aging is responsible for the changes observed.     

Patterns of spatio-temporal correlations in the neural activity of the cat motor cortex during trained forelimb movements

In order to study how neurons in the primary motor cortex (MI) are dynamically linked together during skilled movement, we recorded simultaneously from many cortical neurons in cats trained to perform a reaching and retrieval task using their forelimbs. Analysis of task-related spike activity in the MI of the hemisphere contralateral to the reaching forelimb (in identified forelimb or hindlimb representations) recorded through chronically implanted microwires, was followed by pairwise evaluation of temporally correlated activity in these neurons during task performance using shuffle corrected cross-correlograms. Over many months of recording, a variety of task-related modulations of neural activities were observed in individual efferent zones. Positively correlated activity (mainly narrow peaks at zero or short latencies) was seen during task performance frequently between neurons recorded within the forelimb representation of MI, rarely within the hindlimb area of MI, and never between forelimb and hindlimb areas. Correlated activity was frequently observed between neurons with different patterns of task-related activity or preferential activity during different task elements (reaching, feeding, etc.), and located in efferent zones with dissimilar representation as defined by intracortical microstimulation. The observed synchronization of action potentials among selected but functionally varied groups of MI neurons possibly reflects dynamic recruitment of network connections between efferent zones during skilled movement.     

Role of Acetylcholine in the Modulation of Activity of Neocortical Neurons in Awake Animals Performing an Instrumental Conditioned Reflex

We studied modulatory effects of the cholinergic system on the activity of sensorimotor cortex neurons related to realization of an instrumental conditioned placing reflex. Experiments were carried out on awake cats; multibarrel glass microelectrodes were used for extracellular recording of impulse activity of neurons in the sensorimotor cortex and iontophoretic application of synaptically active agents within the recording region. The background and reflex-related activity was recorded in the course of realization of conditioned movements, and then changes of spiking induced by applications of the testing substances were examined. Applications of acetylcholine and carbachol resulted in increases in the intensity of impulse reactions of neocortical neurons evoked by presentation of an acoustic signal and in simultaneous shortening of the response latencies. An agonist of muscarinic receptors, pylocarpine, exerted a similar effect on the evoked activity of sensorimotor cortex neurons. Blockers of muscarinic receptors, atropine and scopolamine, vice versa, sharply suppressed impulse reactions of cortical neurons to afferent stimulation and simultaneously increased latencies of these responses. Applications of an agonist of nicotinic receptors, nicotine, was accompanied by suppression of impulse neuronal responses, an increase in the latency of spike reactions to presentation of a sound signal, and a corresponding increase in the latency of a conditioned motor reaction. In contrast, application of an antagonist of nicotinic receptors, tubocurarine, significantly intensified neuronal spike responses and shortened their latency. The mechanisms underlying the effects of antagonists of membrane muscarinic and nicotinic cholinoreceptors and the role of activation of these receptors in the modulation of activity of pyramidal and non-pyramidal neocortical neurons related to realization of the instrumental motor reflex are discussed.     

Thermostimulation-Induced Modulation of Conditioned Reflex Movement-Related Reactions of Cortical Neurons of the Cat

In awake cats trained to perform a food-procuring conditioned operant reflex (placing movement), we studied impulse reactions of 86 neurons of the motor cortex (field 4) related to realization of the above movements. As conditioning stimuli (CS) initiating the reflex, we used either non-noxious electrocutaneous stimulation (ECS) of the contralateral forelimb or an acoustic stimulus (sound click). Impulsation of cortical neurons was recorded under conditions of (i) isolated presentation of the CS (control), (ii) presentation of the CS (either ECS or acoustic stimulus) combined with thermostimulation (heating with a miniature electric bulb) of the skin of the working forelimb, and (iii) the same, but with stimulation of the resting forelimb. When we recorded spike activity of neurons within the projection motor zone of the resting limb subjected to ESC, alternating thermostimulation of both forelimbs resulted in considerable intensification and an increase in the duration of neuronal responses, especially in cases where thermostimulation was applied to the working limb ipsilateral to the recording site (a two- to threefold increase). When spike reactions were recorded within the motor cortex of the working forelimb, thermostimulation resulted in a considerable increase in the intensity of these reactions and a decrease in their latency, but only when such stimulation was applied to the working forelimb. Thermostimulation of the resting (ipsilateral, subjected to ESC) limb evoked opposite effects (the intensity of neuronal reactions dropped). In both situations, placing movements remained within the control limits. When sound click was used as a distant CS, thermostimulation of the working limb enhanced neuronal responses, increased their duration by 50-100%, and also increased the time of forestalling of the movement initiation by spike neuronal reactions. Thermostimulation of the resting forelimb in this situation also suppressed neuronal reactions. We conclude that foreign stimulations directed toward modifications of the receptor model of the operant reflex experimental situation formed in the animal result in a decrease in the intensity of the spike responses of field-4 neurons and prolongation of the latencies of these responses, while stimulations promoting the inflow of afferent information to the cortical projection of the working limb evoke opposite effects, an increase in the intensity of neuronal spike responses and a decrease in their latencies.     

Effect of Microiontophoretic Application of Modulators of Noradrenergic Transmission on Impulse Activity of Neurons of the Cat Motor Cortex during Performance of the Reflex to a Complex of Stimuli

On unanesthetized cats, we studied the effects of selective modulators of noradrenergic transmission on the activity of neurons of the cerebral motor cortex in the course of realization of an operant motor reflex to presentation of a complex of stimuli. These modulators were applied using microiontophoretic injections into sites of recording of impulse activity of cortical neurons within the zone of projection of the “working” forelimb. Applications of the α1 receptor agonist Mesaton resulted in significant suppression of background impulsation of the neurons and spiking within the interstimulus interval immediately during realization of the movement. Under the action of the α2 blocker yohimbine, opposite effects were observed. The activity of neurons increased within both background period and other examined time intervals. Mechanisms of the effects of modulation of noradrenergic transmission in the sensorimotor cortex are discussed.     

Unit responses of the first and second somatosensory cortical areas to peripheral,thalamic, and cortical stimulation

Unit responses of the first (SI) somatosensory area of the cortex to stimulation of the second somatosensory area (SII), the ventral posterior thalamic nucleus, and the contralateral forelimb, and also unit responses in SII evoked by stimulation of SI, the ventral posterior thalamic nucleus, and the contralateral forelimb were investigated in experiments on cats immobilized with D-tubocurarine or Myo-Relaxin (succinylcholine). The results showed a substantially higher percentage of neurons in SII than in SI which responded to an afferent stimulus by excitation brought about through two or more synaptic relays in the cortex. In response to cortical stimulation antidromic and orthodromic responses appeared in SI and SII neurons, confirming the presence of two-way cortico-cortical connections. In both SI and SII intracellular recording revealed in most cases PSPs of similar character and intensity, evoked by stimulation of the cortex and nucleus in the same neuron. Latent periods of orthodromic spike responses to stimulation of nucleus and cortex in 50.5% of SI neurons and 37.1% of SII neurons differed by less than 1.0 msec. In 19.6% of SI and 41.4% of SII neurons the latent period of response to cortical stimulation was 1.6–4.7 msec shorter than the latent period of the response evoked in the same neuron by stimulation of the nucleus. It is concluded from these results that impulses from SI play an important role in the afferent activation of SII neurons.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 8, No. 4, pp. 351–357, July–August, 1976.     

Multiple Representations of the Body and Input-Output Relationships in the Agranular and Granular Cortex of the Chronic Awake Guinea Pig

The organization of somatosensory input and the input-output relationships in regions of the agranular frontal cortex (AGr) and granular parietal cortex (Gr) were examined in the chronic awake guinea pig, using the combined technique of single-unit recording and intracortical microstimulation (ICMS). AGr, which was cytoarchitectonically subdivided into medial (AGrm) and lateral (AGrl) parts, also can be characterized on a functional basis. AGrl contains the head, forelimb, and most hindlimb representations; only a small number of hindlimb neurons are confined in AGrm. Different distributions of submodalities exist in AGr and Gr: AGr receives predominantly deep input (with the exception of the vibrissa region, which receives cutaneous input), whereas neurons of Gr respond almost exclusively to cutaneous input. The cutaneous or deep receptive field (RF) of each neuron was determined by natural peripheral stimulation. All studied neurons were activated by small RFs, with the exception of lip, nose, pinna, and limb units of lateral Gr (Grl), for which the RFs were larger.

Microelectrode mapping experiments revealed the existence of three spatially separate, incomplete body maps in which somatosensory and motor representations overlap. One body map, with limbs medially and head rostrolaterally, is contained in AGr. A second map, comparable to the first somatosensory cortex (SI) of other mammals, is found in Gr, with hindlimb, trunk, forelimb, and head representations in an orderly mediolateral sequence. An unresponsive zone separates the head area from the forelimb region. A third map, with the forelimb rostrally and the hindlimb caudally, lies adjacent and lateral to the SI head area. This limb representation, which is characterized by an upright and small size compared to that found in SI, can be considered to be part of the second somatosensory cortex (SII). A distinct head representation was not recognized as properly belonging to SII, but the evidence that neurons of the SI head region respond to stimulation of large RFs located in lips, nose, and pinna leads us to hypothesize that the SII face area overlaps that of SI to some extent, or, alternatively, that the two areas are strictly contiguous and the limits are ambiguous, making them difficult to distinguish.

The input-output relationships were based on the results of RF mapping and ICMS in the same electrode penetration. The intrinsic specific interconnections of cortical neurons whose afferent input and motor output is related to identical body regions show a considerable degree of refinement. The input-output correspondence is especially pronounced for neurons with small RFs. This study confirms and extends similar data recently reported for other rodents.     

Transformation of the afferent tactile signal into a motor command in the cat motor cortex

Activity of 112 neurons of the precruciate motor cortex in cats was studied during a forelimb placing reaction to tactile stimulation of its distal parts. The latent period of response of the limb to tactile stimulation was: for flexors of the elbow (biceps brachii) 30–40 msec, for the earliest reponses of cortical motor neurons about 20 msec. The biceps response was observed 5–10 msec after the end of stimulation of the cortex with a series of pulses lasting 25 msec. Two types of excitatory responses of the neurons were identified: responses of sensory type observed to each tactile stimulation of the limb and independent of the presence or absence of motion, and responses of motor type, which developed parallel with the motor response of the limb and were not observed in the absence of motion. The minimal latent period of the responses of motor type was equal to the latent period of the sensory responses to tactile stimulation (20±10 msec). Stimulation of the cortex through the recording microelectrode at the site of derivation of unit activity, which increased during active flexion of the forelimb at the elbow (11 stimuli at intervals of 2.5 msec, current not exceeding 25 µA), in 70% of cases evoked an electrical response in the flexor muscle of the elbow.M. V. Lomonosov Moscow State University. Translated from Neirofiziologiya, Vol. 9, No. 2, pp. 115–123, March–April, 1977.     

Areal distributions of cortical neurons projecting to different levels of the caudal brain stem and spinal cord in rats

Distributions of corticospinal and corticobulbar neurons were revealed by tetramethylbenzidine (TMB) processing after injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) into the cervical or lumbar enlargements of the spinal cord, or medullary or pontine levels of the brain stem. Sections reacted for cytochrome oxidase (CO) allowed patterns of labeled neurons to be related to the details of the body surface map in the first somatosensory cortical area (SI). The results indicate that a number of cortical areas project to these subcortical levels: (1) Projection neurons in granular SI formed a clear somatotopic pattern. The hindpaw region projected to the lumbar enlargement, the forepaw region to the cervical enlargement, the whisker pad field to the lower medulla, and the more rostral face region to more rostral brain stem levels. (2) Each zone of labeled neurons in SI extended into adjacent dysgranular somatosensory cortex, forming a second somatotopic pattern of projection neurons. (3) A somatotopic pattern of projection neurons in primary motor cortex (MI) paralleled SI in mediolateral sequence corresponding to the hindlimb, forelimb, and face. (4) A weak somatotopic pattern of projection neurons was suggested in medial agranular cortex (Agm), indicating a premotor field with a rostromedial-to-caudolateral representation of hindlimb, forelimb, and face. (5) A somatotopic pattern of projection neurons representing the foot to face in a mediolateral sequence was observed in medial parietal cortex (PM) located between SI and area 17. (6) In the second somatosensory cortical area (SII), neurons projecting to the brain stem were immediately adjacent caudolaterally to the barrel field of SI, whereas neurons projecting to the upper spinal cord were more lateral. No projection neurons in this region were labeled by the injections in the lower spinal cord. (7) Other foci of projection neurons for the face and forelimb were located rostral to SII, providing evidence for a parietal ventral area (PV) in perirhinal cortex (PR) lateral to SI, and in cortex between SII and PM. None of these regions, which may be higher-order somatosensory areas, contained labeled neurons after injections in the lower spinal cord. Thus, more cortical fields directly influence brain stem and spinal cord levels related to sensory and motor functions of the face and forepaw than the hindlimb. The termination patterns of corticospinal and corticobulbar projections were studied in other rats with injections of WGA:HRP in SI.(ABSTRACT TRUNCATED AT 400 WORDS)     

A computer model of generation of the motor cortex neuron processes seen in the course of execution of instrumental movement

A computer model of neuronal processes in the motor cortex column is presented. The model is consisted of two pyramidal cell layers with two groups of inhibitory interneurons, selectively controlling pyramidal cell soma and dendrite, in each. Active Na, Ca and K conductances are included in the model of a single neuron. Horizontal excitatory connections between pyramidal cells in the upper layer are largely of NMDA-receptor type, that in the lower layer--of non-NMDA-type. All inhibitory synapses are of GABA(A)-type. The model reproduces the main phenomenon observed in the motor cortex during the execution of conditioned movements. Consequent to an early excitation the upper layer pyramidal cells generate a late NMDA-dependent reflexive response to afferent conditional stimulation, which as in a real case is diminished by GABA(A)-type synaptic inhibition and afferent stimulus strength increase. The characteristic inverse relation between the late response manifestation and the stimulus strength observed in the real cortex can be reproduced in the model only if NMDA-glutamate receptors were preferentially localized in the terminals of pyramidal cell backward collaterals, not in the terminals of the afferent fibers on pyramidal neurons. The intended component of motor cortex neuronal activity is generated in NMDA-independent manner by the pyramidal cells of lower layer. The slow time coarse of intended component as compared with short duration of AMPA epsp's is due to a consecutive relay-race--like activation of pyramidal neurons with different dendrit-to-soma ratio.     

Areal Distributions of Cortical Neurons Projecting to Different Levels of the Caudal Brain Stem and Spinal Cord in Rats

Distributions of corticospinal and corticobulbar neurons were revealed by tetramethylbenzidine (TMB) processing after injections of wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) into the cervical or lumbar enlargements of the spinal cord, or medullary or pontine levels of the brain stem. Sections reacted for cytochrome oxidase (CO) allowed patterns of labeled neurons to be related to the details of the body surface map in the first somatosensory cortical area (SI). The results indicate that a number of cortical areas project to these subcortical levels: (1) Projection neurons in granular SI formed a clear somatotopic pattern. The hindpaw region projected to the lumbar enlargement, the forepaw region to the cervical enlargement, the whisker pad field to the lower medulla, and the more rostral face region to more rostral brain stem levels. (2) Each zone of labeled neurons in SI extended into adjacent dysgranular somatosensory cortex, forming a second somatotopic pattern of projection neurons. (3) A somatotopic pattern of projection neurons in primary motor cortex (MI) paralleled SI in mediolateral sequence corresponding to the hindlimb, forelimb, and face. (4) A weak somatotopic pattern of projection neurons was suggested in medial agranular cortex (Ag_m), indicating a premotor field with a rostromedial-to-caudolateral representation of hindlimb, forelimb, and face. (5) A somatotopic pattern of projection neurons representing the foot to face in a mediolateral sequence was observed in medial parietal cortex (PM) located between SI and area 17. (6) In the second somatosensory cortical area (SII), neurons projecting to the brain stem were immediately adjacent caudolaterally to the barrel field of SI, whereas neurons projecting to the upper spinal cord were more lateral. No projection neurons in this region were labeled by the injections in the lower spinal cord. (7) Other foci of projection neurons for the face and forelimb were located rostral to SII, providing evidence for a parietal ventral area (PV) in perirhinal cortex (PR) lateral to SI, and in cortex between SII and PM. None of these regions, which may be higher-order somatosensory areas, contained labeled neurons after injections in the lower spinal cord. Thus, more cortical fields directly influence brain stem and spinal cord levels related to sensory and motor functions of the face and forepaw than the hindlimb.

The termination patterns of corticospinal and corticobulbar projections were studied in other rats with injections of WGA:HRP in SI. Injections in lateral SI representing the face produced dense terminal label in the contralateral trigeminal complex. Injections in cortex devoted to the forelimb and forepaw labeled the contralateral cuneate nucleus and parts of the dorsal horn of the spinal cord. The cortical injections also demonstrated interconnections of parts of SI with some of the other regions of cortex with projections to the spinal cord, and provided further evidence for the existence of PV in rats.     

Comparison of the Connectional Properties of the Two Forelimb Areas of the Rat Sensorimotor Cortex: Support for the Presence of a Premotor or Supplementary Motor Cortical Area

The existence of multiple motor cortical areas that differ in some of their properties is well known in primates, but is less clear in the rat. The present study addressed this question from the point of view of connectional properties by comparing the afferent and efferent projections of the caudal forelimb area (CFA), considered to be the equivalent of the forelimb area of the primary motor cortex (MI), and a second forelimb motor representation, the rostral forelimb area (RFA). As a result of various tracing experiments (including double labeling), it was observed that CFA and RFA had reciprocal corticocortical connections characterized by preferential, asymmetrical, laminar distribution, indicating that RFA may occupy a different hierarchical level than CFA, according to criteria previously discussed in the visual cortex of primates. Furthermore, it was found that RFA, but not CFA, exhibited dense reciprocal connections with the insular cortex. With respect to their efferent projection to the basal ganglia, it was observed that CFA projected very densely to the lateral portion of the ipsilateral caudate putamen, whereas the contralateral projection was sparse and more restricted. The ipsilateral projection originating from RFA was slightly less dense than that from CFA, but it covered a larger portion of the caudate putamen (in the medial direction); the contralateral projection from RFA to the caudate putamen was of the same density and extent as the ipsilateral projection. The reciprocal thalamocortical and corticothalamic connections of RFA and CFA differed from each other in the sense that CFA was mainly interconnected with the ventrolateral thalamic nucleus, while RFA was mainly connected with the ventromedial thalamic nucleus. Altogether, these connectional differences, compared with the pattern of organization of the motor cortical areas in primates, suggest that RFA in the rat may well be an equivalent of the premotor or supplementary motor area. In contrast to the corticocortical, corticostriatal, and thalamocortical connections, RFA and CFA showed similar efferent projections to the subthalamic nucleus, substantia nigra, red nucleus, tectum, pontine nuclei, inferior olive, and spinal cord.     

Regulation of afferent transmission in the feeding circuitry of Aplysia

Although feeding in Aplysia is mediated by a central pattern generator (CPG), the activity of this CPG is modified by afferent input. To determine how afferent activity produces the widespread changes in motor programs that are necessary if behavior is to be modified, we have studied two classes of feeding sensory neurons. We have shown that afferent-induced changes in activity are widespread because sensory neurons make a number of synaptic connections. For example, sensory neurons make monosynaptic excitatory connections with feeding motor neurons. Sensori-motor transmission is, however, regulated so that changes in the periphery do not disrupt ongoing activity. This results from the fact that sensory neurons are also electrically coupled to feeding interneurons. During motor programs sensory neurons are, therefore, rhythmically depolarized via central input. These changes in membrane potential profoundly affect sensori-motor transmission. For example, changes in membrane potential alter spike propagation in sensory neurons so that spikes are only actively transmitted to particular output regions when it is behaviorally appropriate. To summarize, afferent activity alters motor output because sensory neurons make direct contact with motor neurons. Sensori-motor transmission is, however, centrally regulated so that changes in the periphery alter motor programs in a phase-dependent manner.     

The role of the motor cortex in rearrangement of the innate movement coordination in the dog

In chronical experiments in dogs the pattern of shoulder muscle recruitment was examined during the forelimb flexion by which the animal lifted and held a cup of food during eating. At the early stage of the instrumental reaction learning the forelimb lifting was performed with the anticipatory deviation of the head in up direction, when the head bent down to the foodwell the lifted forelimb lowered. Simultaneous holding of the flexed forelimb and lowered head providing food reinforcement was achieved only by learning. It was found that the forelimb lifting in the innate coordination in untrained dogs was performed with activation of m. deltoideus and m. teres major, whereas m. teres minor was active whilst the dog was standing but the muscle activity was abolished before the limb lifting. In the course of learning m. teres minor activity was changed into opposite one. In the learned coordination the limb lifting was accompanied by the activation of all three shoulder flexors. The lesion of the motor cortex in the area of the "working" forelimb, but not in other areas led to disturbance of the learned coordination and the novel pattern of the shoulder muscle activity. The data obtained led to the following conclusion: the rearrangement of the innate coordination is connected with the formation of the novel way of the forelimb lifting which pattern of muscle recruitment is provided by the motor cortex.     

Impulse activity of the bulbar cardiovascular neurons during myocardial ischemia and stimulation of the cortical sensorimotor zone

In acute experiments on cats performed under nembutal anesthesia the stimulation of sensorimotor zone in cerebral hemisphere cortex changed the impulse activity of interneurons of bulbar cardiovascular centre and not of the afferent neurons. The analysis of the activity of afferent neurons and interneurons has shown a decrease in coordination between the reaction of these cells to the development of ischemic myocardial lesions during the cortex stimulation. In these conditions bulbar cardiovascular neurons could both increase and decrease the impulse activity. These changes seem to be the reason for the growing incidence of idioventricular ischemic arrhythmias during cortical stimulation.     

Acquisition of conditioned facial reflexes in the cat: cortical control of different facial movements

The motor cortex plays a role in determining which of three different facial movements is acquired in Pavlovian conditioning experiments. Three separate facial reflexes can be distinguished by recording electromyographic activity from the orbicularis oculi (eye blink) and levator orii (nose twitch) muscles. One in a pure eye blink; a second is a nose twitch; the third is a compound eye blink and nose twitch. Which of these movements is elicited by a click (conditioned stimulus) following associative conditioning is reflected by the pattern of unit activity elicited by the click at the motor cortex. Activity is enhanced, after conditioning, in those units that project polysynaptically to the specific muscles performing the learned movement. This enhancement of activity is, in turn, relatable to an enhanced electrical excitability of the involved neurons. Analogous changes in the excitability of neurons of the motor cortex to applied currents can be produced by local application of cholinergic agents. Iontophoresis of acetylcholine, aceclidine (a cholinomimetic drug), or intracellularly applied cyclic GMP produces changes in single neuron membrane resistance that increase neuronal excitability. The units of the motor cortex that respond preferentially to these agents and to the click conditioned stimuli with short latencies have been identified as pyramidal cells of layer V. The axons of these neurons form the pyramidal tract, a pathway characterized as serving voluntary movement. It appears that this system supports rapid transmission and processing of auditory-motor information used to perform learned movements adaptively, selectively, and discriminatively.     

Specification of synaptic connections between sensory and motor neurons in the developing spinal cord

Experimental studies of mechanisms underlying the specification of synaptic connections in the monosynaptic stretch reflex of frogs and chicks are described. Sensory neurons innervating the triceps brachii muscles of bullfrogs are born throughout the period of sensory neurogenesis and do not appear to be related clonally. Instead, the peripheral targets of these sensory neurons play a major role in determining their central connections with motoneurons. Developing thoracic sensory neurons made to project to novel targets in the forelimb project into the brachial spinal cord, which they normally never do. Moreover, these foreign sensory neurons make monosynaptic excitatory connections with the now functionally appropriate brachial motoneurons. Normal patterns of neuronal activity are not necessary for the formation of specific central connections. Neuromuscular blockade of developing chick embryos with curare during the period of synaptogenesis still results in the formation of correct sensory-motor connections. Competitive interactions among the afferent fibers also do not seem to be important in this process. When the number of sensory neurons projecting to the forelimb is drastically reduced during development, each afferent still makes central connections of the same strength and specificity as normal. These results are discussed with reference to the development of retinal ganglion cells and their projections to the brain. Although many aspects of the two systems are similar, patterned neural activity appears to play a much more important role in the development of the visual pathway than in the spinal reflex pathway described here.     

Involvement of rabbit motor cortex neurons in the instrumental behavior before and after chronic ethanol consumption: a comparison with the limbic cortex

The effect of acute administration is significantly more prominent in the limbic (cingulate) than in the motor cortex [9]. We proposed that the limbic cortex is more sensitive also to chronic ethanol treatment (CET). It was shown that morphology as well as neuronal activity of the limbic cortex changed greatly after the CET [7]. The missing link of testing the above proposition was a comparison of the obtained data with the results of the experimental study of CET influence on the motor cortex. Morphology of the anterolateral motor cortex and activity of its neurons in the instrumental food-acquisition behavior were studied in 6 male rabbits after CET (9 months). It was found that the limbic cortex was modified morphologically and functionally to a significantly greater extent than the motor cortex. We consider the fact that in the limbic cortex of a healthy individual there are many neurons, which for a while cease their discharges after the acute ethanol administration, to be among the most important reasons for this difference. Such-like repeated activity interruptions in the course of CET impair the performance of the systems incorporating these neurons. In such a way ethanol prevents all neurons, especially the mentioned ones, from receiving adequate metabolic supply that is necessary for their survival and functioning.     

A Comparison of the Ipsilateral Cortical Projections to the Dorsal and Ventral Subdivisions of the Macaque Premotor Cortex

The cortical connections of the dorsal (PMd) and ventral (PMv) subdivisions of the premotor area (PM, lateral area 6) were studied in four monkeys (Macaca fascicularis) through the use of retrograde tracers. In two animals, tracer was injected ventral to the arcuate sulcus (PMv), in a region from which forelimb movements could be elicited by intracortical microstimulation (ICMS). Tracer injections dorsal to the arcuate sulcus (PMd) were made in two locations. In one animal, tracer was injected caudal to the genu of the arcuate sulcus (in caudal PMd [cPMd], where ICMS was effective in eliciting forelimb movements); in another animal, it was injected rostral to the genu of the arcuate sulcus (in rostral PMd [rPMd], where ICMS was ineffective in eliciting movements). Retrogradely labeled neurons were counted in the ipsilateral hemisphere and located in cytoarchitectonically identified areas of the frontal and parietal lobes. Although both PMv and PMd were found to receive inputs from other motor areas, the prefrontal cortex, and the parietal cortex, there were differences in the topography and the relative strength of projections from these areas.

There were few common inputs to PMv and PMd; only the supplementary eye fields projected to all three areas studied. Interconnections within PMd or PMv appeared to link hindlimb and forelimb representations, and forelimb and face representations; however, connections between PMd and PMv were sparse. Areas cPMd and PMv were found to receive inputs from other motor areas—the primary motor area, the supplementary motor area, and the cingulate motor area—but the topography and strength of projections from these areas varied. Area rPMd was found to receive sparse inputs, if any, from these motor areas. The frontal eye field (area 8a) was found to project to PMv and rPMd, and area 46 was labeled substantially only from rPMd. Parietal projections to PMv were found to originate from a variety of somatosensory and visual areas, including the second somatosensory cortex and related areas in the parietal operculum of the lateral sulcus, as well as areas 5, 7a, and 7b, and the anterior intraparietal area. By contrast, projections to cPMd arose only from area 5. Visual areas 7m and the medial intraparietal area were labeled from rPMd. Relatively more parietal neurons were labeled after tracer injections in PMv than in PMd. Thus, PMv and PMd appear to be parts of separate, parallel networks for movement control.     

Disruption of acquired head and paw movement coordination after unilateral lesioning of the motor cortex in dogs (a kinematic analysis)

Dogs were trained to remove a cup with meat to the head bent down to the feeder and hold the limb flexed during eating. At the early stage of learning, the stable innate head-forelimb coordination characteristic for untrained animals was manifest. The forelimb flexion was accompanied by anticipatory lifting of the bent head, and the following bending of the head led to an extension of the flexed forelimb. The opposite coordination, i.e., the lifting and holding of the forelimb when the head is bent down, was achieved only by training. The lesion of the motor cortex contralateral to the working forelimb in the trained dogs led to a prolonged disturbance of the simultaneous holding of the flexed forelimb and the head bent down. The lesion of the motor cortex did not affect the individual movements but disturbed their coordination. In the operated dogs the innate relationships between the head and forelimb movement recovered. The results support the previous finding that the lesion of the motor cortex led to recovery of the innate coordination transformed in the process of learning.