Activity of the neostriatum neurons in monkeys during operant behavior

Monkeys (Macaca nemestrina) were trained to perform an operant task which included the following stages: bimanual movement to a visible target in response to a start signal, selection of the left or right operating hand according to the visual conditioning signal, careful movement of the selected hand, movements of fingers of the same hand, and taking of a reward. The spike activity was recorded simultaneously from five or six neurons in the caudate nucleus (Cd) head. Most of the Cd neurons were activated during the taking of a reward, the analysis of conditioning signals, and visually controlled movements. It has been found that the same neuron of the Cd may be activated in different ways during different components of the behavioral program, depending on the complexity of the conditioning signals and the correct versus incorrect selection of an operating hand. It is concluded that the Cd neurons have no functional specialization and that they take part in mechanisms preceding a movement or accompanying it, but not correlating with the form of its realization.Translated from Neirofiziologiya, Vol. 25, No. 2, pp. 132–140, March–April, 1993.     

Self-Initiated Behavioral Act-Related Neuronal Activity in the Region of the Raphe Nuclei of the Cat

In experiments on awake cats, we recorded the activity of 79 putative serotonergic (STE) neurons localized within the region of the brainstem dorsal and anterior central raphe nuclei. The animals were trained to perform a self-initiated (voluntary) movement, to press a pedal by the forelimb; an additional limitation was to perform the movement not earlier than after a definite time interval. Changes in the activity of STE neurons related to the preparation for and performance of the movement and reactions to presentation of a feedback conditioning signal preceding the reward and receipt of the food reward were most clearly manifested. More than 50% of the units changed their activity before the movement initiation. Most neurons responded to presentation of a positive conditioning signal by phasic activation, while a negative signal informing them of the absence of the reward evoked considerably weaker reactions. We hypothesize that reactions of STE neurons forestalling the movement initiation can provide activation of the neocortex necessary for the movement performance within a preset time interval. Activating and inhibitory reactions observed within the period of expectation of a feedback conditioning signal and developing after presentation of this signal can be related to a noticeable role of the STE system in the formation of memory engrams and development of emotional states.     

Spatiotemporal organization of unit activity in the frog tectum and its modification by visual stimuli

Unit activity was recorded in the tectum of curarized frogs during presentation of various visual stimuli (on- and off-responses to diffuse illumination and movement of an object of recognizable shape). It was shown that different types of stimulation lead to the organization of a different distribution of unit activity in the tectum, in the form of excitatory-inhibitory neuronal mosaics; inhibitory responses, limiting and exacerbating the excitatory responses of other neurons, predominate. The differences observed in the spatiotemporal characteristics of the neuronal mosaics under the influence of different stimuli may be evidence of specificity of coding of visual impulses carrying different information from the retina in the tectum.     

Collective reactions of neostriatum (putamen) neurons to alternative behaviour in monkeys

Significant differences were found in collective unit activity of 6 neostriatum neurons associated with performance of either left- or right-sided task by monkeys: at the moment of decision-making regarding the direction of the movement, and after its completion, irrespective of correctness of the choice. The differences at the moment of the programme completion were significantly greater than at the moment of the decision-making.     

Unit activity of the prefrontal cortex during delayed alternation performance in monkey.

Unit activity in the dorsolateral regions of the prefrontal cortex (s. principalis) of the monkey was analysed by multineuronal recording technique. The sequence during delayed-response alternation included anticipatory stimulus, non-specific expectancy, conditioned cue, delay, trigger stimulus and alternation performance. Food reward completed the sequence. Unit activity in s. principalis was found to be involved in such learning processes as integration of behavioural acts into an accomplished programme and storage of traces in short-term memory. The latter can be observed in successive involvement of neuronal populations during a 10 sec delay as well as in rearrangements of unit activity, the maximum of which shifts to the latest part of the delay period. Unit activity in s. principalis reflects the level of correct response, corresponding correlates of which depend on the level of attention. The dorsolateral prefrontal cortex has structural and functional characteristics for short-term storage of external signals in its neuronal nets according to the level of attention.     

Memory for spatial and object-specific cues in food-storing and non-storing birds

Two storer/non-storer pairs of species, marsh tit (Parus palustris)/blue tit (P. caeruleus) and jay (Garrulus glandarius)/jackdaw (Corvus monedula) were compared on a one-trial associative memory task. In phase I of a trial birds searched for a reward in one of four feeders which differed in their trial-unique spatial location and object-specific cues. Following a retention interval, the birds had to return to the same feeder to obtain a further reward. In control trials the array of feeders was unaltered, whilst in dissociation tests it was transformed to separate spatial location and object-specific cues.In control trials there was no difference in performance between species. In dissociation tests, the two storing species went first to the correct spatial location and second to the correct object-specific cues, whereas the two non-storing species went first with equal probability to the correct spatial and local object cues.Monocular occlusion was used to investigate differences between the two eye-systems. In control trials there was no effect of occlusion. In dissociation trials, all 4 species preferentially returned to the feeder with the correct object-specific cue when the left eye had been covered in phase I and to the feeder in the correct spatial position when the right eye had been covered in phase I.These results suggest that (a) food-storing birds differ from non-storers in responding preferentially to spatial information and (b) in storers and non-storers the right eye system shows a preference for object-specific cues and the left eye system for spatial cues.     

Reorganization of Composition of Striatal Neurons Correlating with Behavior in the Monkey Macaca nemestrina

The available experimental data do not provide a sufficiently complete picture of the neuronal activity connected with some action of the animal, as they have been obtained under different experimental protocols and as a result of study of different cells. The present work was aimed at studying activity of the same neuron group at different moments of animal's behavior. A monkey (Macaca nemestrina) was taught to perform a behavioral program consisting of several functional heterogeneous actions. The impulsive activity of striatal neurons was recorded in the central region of putamen with coordinates A 16.5, L7, and H 8–10 [18]. The activity of each neuron was recorded during 13 consecutive stages of the same behavioral task. As a whole, in 59 putamen neurons, 767 fragments of neuronal activity were studied. It was shown that the same neurons could be involved at different behavioral stages when the animal performed different actions. At individual stages, the number of neurons common with other behavioral stages reached 70–80% of all reactive cells at the stage. The number of the neurons common within the rest of 12 stages was determined for every program stage. The number of such common neurons established in the experiment was in 142 out of 156 cases higher than their number that could be expected on the basis of statistical relations. The data obtained indicate that the reorganization in composition of behavior-reactive cells at every behavioral stage occurs mainly by using the same neurons but not only the neurons that are specialized for the given action. The polymodality of individual striatal neurons is unlikely to be connected with that they have several functions, but results from that the same neuron can be a constituent of neuronal mosaics of different configurations corresponding to different behavioral moments.     

Differentiating activity of monkey putamen neurons during performance of the alternative spatial choice

Single unit activity was recorded in monkeys in three putamen zones learned a bimanual operant activity during performance of the task of alternative spatial choice. The neuronal reactions were specially analyzed by the criteria as follows: a) differentiation of the side of reward (differentiating--non-differentiating reactions); b) character of reaction by duration (tonic-phasic); c) laterality (contra- and ipsilateral reactions as related to hemisphere); d) frequency of background activity. It was shown that differentiating cell activity, especially their tonic part and in still greater degree contra-lateral tonic reactions most closely correlate with behavioral aspects of the program. The assumption that differentiating activity, unlike non-differentiating one, is the reflection of not only morphological and neurochemical characteristic features of nervous elements of putamen but of its functional uniformity in relation to external determinants of behavior, was put forward.     

Coding the differently directed behavioral actions by the putamen neurons of the monkey brain

Spike activity related to the problem of alternative choice of behavioral actions was recorded in the putamen of the monkey brain. The patterns of low and high activities were identified. Each neuron during different behaviour actions could generate any kind of patterns. The differences between neuronal compositions with patterns of high activity, at the left and right direction of the task, were obtained during decision making about the movement direction, and also at the end of the movement. Distinctions between neuronal compositions with patterns of low activity at this time, on the contrary, diminished. The neuronal compositions with patterns of low activity were much more before the conditioned signal, when the animal did not yet know the task, and at the end of the program when the problem was already solved. The data obtained show that the putamen units control different directions of actions by a multilevel address coding, mainly through reorganizing the neuronal compositions with patterns of different level activity.     

Duplicated rDNA sequences of variable lengths flanking the short type I insertions in the rDNA of Drosophila melanogaster.

We describe cloned segments of rDNA that contain short type I insertions of differing lengths. These insertions represent a coterminal subset of sequences from the right hand side of the major 5kb type I insertion. Three of these shorter insertions are flanked on both sides by a short sequence present as a single copy in uninterrupted rDNA units. The duplicated segment is 7, 14 and 15 nucleotides in the different clones. In this respect, the insertions differ from the 5kb type I insertion, where the corresponding sequence is found only at the right hand junction and where at the left hand side there is a deletion of 9 nucleotides of rDNA (Roiha et al.,1981). One clone is unusual in that it contains two type I insertions, one of which is flanked by a 14 nucleotide repeat. The left hand junction of the second insertion occurs 380 nucleotides downstream in the rDNA unit from the first. It has an identical right hand junction to the other elements and the 380 nucleotide rDNA sequence is repeated on both sides of the insertion. We discuss the variety of sequence rearrangements of the rDNA which flank type I insertions.     

Neuronal activity in the primate supplementary motor area and the primary motor cortex in relation to spatio-temporal bimanual coordination

Single neuronal activity was recorded from the supplementary motor area (SMA-proper and pre-SMA) and primary motor cortex (M1) in two Macaca fascicularis trained to perform a delayed conditional sequence of coordinated bimanual pull and grasp movements. The behavioural paradigm was designed to distinguish neuronal activity associated with bimanual coordination from that related to a comparable motor sequence but executed unimanually (left or right arm only). The bimanual and unimanual trials were instructed in a random order by a visual cue. Following the cue, there was a waiting period until presentation of a "go-signal", signalling the monkey to perform the instructed movement. A total of 143 task-related neurons were recorded from the SMA (SMA-proper, 62; pre-SMA, 81). Most SMA units (87%) were active in both unimanual contralateral and unimanual ipsilateral trials (bilateral neurons), whereas 9% of units were active only in unimanual contralateral trials and 3% were active only in unimanual ipsilateral trials. Forty-eight per cent of SMA task-related units were classified as bimanual, defined as neurons in which the activity observed in bimanual trials could not be predicted from that associated with unimanual trials when comparing the same events related to the same arm. For direct comparison, 527 neurons were recorded from M1 in the same monkeys performing the same tasks. The comparison showed that M1 contains significantly less bilateral neurons (75%) than the SMA, whereas the reverse was observed for contralateral neurons (22% in M1). The proportion of M1 bimanual cells (53%) was not statistically different from that observed in the SMA. The results suggest that both the SMA and M1 may contribute to the control of sequential bimanual coordinated movements. Interlimb coordination may then take place in a distributed network including at least the SMA and M1, but the contribution of other cortical and subcortical areas such as cingulate motor cortex and basal ganglia remains to be investigated.     

Asymmetric activation of the primary motor cortex during observation of a mirror reflection of a hand

Mirror therapy is an effective technique for pain relief and motor function recovery. It has been demonstrated that magnetic 20-Hz activity is induced in the primary motor cortex (M1) after median nerve stimulation and that the amount of the stimulus-induced 20-Hz activity is decreased when the M1 is activated. In the present study, we investigated how the image or the mirror reflection of a hand holding a pencil modulates the stimulus-induced 20-Hz activity in the M1. Neuromagnetic brain activity was recorded from 13 healthy right-handed subjects while they were either viewing directly their hand holding a pencil or viewing a mirror reflection of their hand holding a pencil. The 20-Hz activity in the left or the right M1 was examined after the right or the left median nerve stimulation, respectively, and the suppression of the stimulus-induced 20-Hz in the M1 by viewing directly one hand holding a pencil or by viewing the mirror image of the hand holding a pencil was assumed to indicate the activation of the M1. The results indicated that the M1 innervating the dominant hand was suppressed either by viewing directly the dominant hand holding a pencil or by viewing the mirror image of the non-dominant hand holding a pencil. On the other hand, the M1 innervating the non-dominant hand was activated by viewing the mirror image of the dominant hand holding a pencil, but was not activated by viewing directly the non-dominant hand holding a pencil. The M1 innervating either the dominant or the non-dominant hand, however, was not activated by viewing the hand on the side ipsilateral to the M1 examined or the mirror image of the hand on the side contralateral to the M1 exaimined. Such activation of the M1 might induce some therapeutic effects of mirror therapy.     

Excitability of the motor cortex ipsilateral to the moving body side depends on spatio-temporal task complexity and hemispheric specialization

Unilateral movements are mainly controlled by the contralateral hemisphere, even though the primary motor cortex ipsilateral (M1(ipsi)) to the moving body side can undergo task-related changes of activity as well. Here we used transcranial magnetic stimulation (TMS) to investigate whether representations of the wrist flexor (FCR) and extensor (ECR) in M1(ipsi) would be modulated when unilateral rhythmical wrist movements were executed in isolation or in the context of a simple or difficult hand-foot coordination pattern, and whether this modulation would differ for the left versus right hemisphere. We found that M1(ipsi) facilitation of the resting ECR and FCR mirrored the activation of the moving wrist such that facilitation was higher when the homologous muscle was activated during the cyclical movement. We showed that this ipsilateral facilitation increased significantly when the wrist movements were performed in the context of demanding hand-foot coordination tasks whereas foot movements alone influenced the hand representation of M1(ipsi) only slightly. Our data revealed a clear hemispheric asymmetry such that MEP responses were significantly larger when elicited in the left M1(ipsi) than in the right. In experiment 2, we tested whether the modulations of M1(ipsi) facilitation, caused by performing different coordination tasks with the left versus right body sides, could be explained by changes in short intracortical inhibition (SICI). We found that SICI was increasingly reduced for a complex coordination pattern as compared to rest, but only in the right M1(ipsi). We argue that our results might reflect the stronger involvement of the left versus right hemisphere in performing demanding motor tasks.     

Unit responses of the clare-bishop cortical association area in the cat to photic stimulation

Electrical activity of single unit in the Clare-Bishop visual association area of the cortex was studied in acute experiments on cats immobilized with Flaxedil and after pretrigeminal sections. The method of extracellular recording of action potentials of single units was used. The experimental results showed that 95.5% of cells responding to visual stimulation responded to movement of a spot of light in the receptive field of the neurons, and 55% of the cells responded selectively to the direction of movement. Some neurons responded to movement of a stimulus only when it entered and left the receptive field. About 85.3% of cells responded to a flashing spot of light, and also to a general change in the intensity of illumination of the receptive field. The receptive field of neurons of the Clare-Bishop area in most cases were in the form of stripes with their long axis horizontal. The results point to the important role of this cortical association area in the central analysis of visual information.L. A. Orbeli Institute of Physiology, Academy of Sciences of the Armenian SSSR, Erevan. Translated from Neirofiziologiya, Vol. 10, No. 1, pp. 22–29, January–February, 1978.     

Comparative analysis of diadochokinetic movements.

Tests of diadochokinesia are an inherent part of a neurological examination. Various quantifying methods have been proposed to increase the objectivity, sensitivity, and reliability of such examinations. The methods used and analyses performed, however, differ substantially between tasks. We used a three-dimensional, ultrasound-based recording device to continuously record joint angles during three diadochokinetic movements, avoiding any external constraints of the movements. Alternate pronation and supination of the forearm, tapping with the whole hand and with the index finger in isolation were analyzed in a sample of 63 healthy control subjects. The most sensitive measure for capturing effects of gender, sex, and active hand was frequency. The right hand was faster than the left in all tasks, tapping performance declined with increasing age, and male subjects were faster than females in forearm diadochokinesia. Other measures that characterize speed of movement such as maximum angular velocities and accelerations did not yield comparable sensitivity in detecting the same statistical effects. However, angular velocity achieved the highest test-retest reliability for forearm diadochokinesia, while frequency was reproduced in the tapping tasks. Additional measures characterizing symmetry of the angular velocity profiles and intraindividual variability were shown to be largely independent of movement speed. Examples in neurological patients showed that the data define a valuable standard against which pathological performance can be precisely evaluated. In addition, the different measures captured dissociable aspects of motor performance that may further help to characterize the deficit and adjust therapy.     

Value representations in the primate striatum during matching behavior

Choosing the most valuable course of action requires knowing the outcomes associated with the available alternatives. The striatum may be important for representing the values of actions. We examined this in monkeys performing an oculomotor choice task. The activity of phasically active neurons (PANs) in the striatum covaried with two classes of information: action-values and chosen-values. Action-value PANs were correlated with value estimates for one of the available actions, and these signals were frequently observed before movement execution. Chosen-value PANs were correlated with the value of the action that had been chosen, and these signals were primarily observed later in the task, immediately before or persistently after movement execution. These populations may serve distinct functions mediated by the striatum: some PANs may participate in choice by encoding the values of the available actions, while other PANs may participate in evaluative updating by encoding the reward value of chosen actions.     

Neuron selection by relative importance for neural decoding of dexterous finger prosthesis control application

Future generations of upper limb prosthesis will have dexterous hand with individual fingers and will be controlled directly by neural signals. Neurons from the primary motor (M1) cortex code for finger movements and provide the source for neural control of dexterous prosthesis. Each neuron's activation can be quantified by the change in firing rate before and after finger movement, and the quantified value is then represented by the neural activity over each trial for the intended movement. Since this neural activity varies with the intended movement, we define the relative importance of each neuron independent of specific intended movements. The relative importance of each neuron is determined by the inter-movement variance of the neural activities for respective intended movements. Neurons are ranked by the relative importance and then a subpopulation of rank-ordered neurons is selected for the neural decoding. The use of the proposed neuron selection method in individual finger movements improved decoding accuracy by 21.5% in the case of decoding with only 5 neurons and by 9.2% in the case of decoding with only 10 neurons. With only 15 highly ranked neurons, a decoding accuracy of 99.5% was achieved. The performance improvement is still maintained when combined movements of two fingers were included though the decoding accuracy fell to 95.7%. Since the proposed neuron selection method can achieve the targeting accuracy of decoding algorithms with less number of input neurons, it can be significant for developing brain–machine interfaces for direct neural control of hand prostheses.     

Direction of movement is encoded in the human primary motor cortex

The present study investigated how direction of hand movement, which is a well-described parameter in cerebral organization of motor control, is incorporated in the somatotopic representation of the manual effector system in the human primary motor cortex (M1). Using functional magnetic resonance imaging (fMRI) and a manual step-tracking task we found that activation patterns related to movement in different directions were spatially disjoint within the representation area of the hand on M1. Foci of activation related to specific movement directions were segregated within the M1 hand area; activation related to direction 0° (right) was located most laterally/superficially, whereas directions 180° (left) and 270° (down) elicited activation more medially within the hand area. Activation related to direction 90° was located between the other directions. Moreover, by investigating differences between activations related to movement along the horizontal (0°+180°) and vertical (90°+270°) axis, we found that activation related to the horizontal axis was located more anterolaterally/dorsally in M1 than for the vertical axis, supporting that activations related to individual movement directions are direction- and not muscle related. Our results of spatially segregated direction-related activations in M1 are in accordance with findings of recent fMRI studies on neural encoding of direction in human M1. Our results thus provide further evidence for a direct link between direction as an organizational principle in sensorimotor transformation and movement execution coded by effector representations in M1.     

Electrical stimulation of the primate lateral habenula suppresses saccadic eye movement through a learning mechanism

The lateral habenula (LHb) is a brain structure which represents negative motivational value. Neurons in the LHb are excited by unpleasant events such as reward omission and aversive stimuli, and transmit these signals to midbrain dopamine neurons which are involved in learning and motivation. However, it remains unclear whether these phasic changes in LHb neuronal activity actually influence animal behavior. To answer this question, we artificially activated the LHb by electrical stimulation while monkeys were performing a visually guided saccade task. In one block of trials, saccades to one fixed direction (e.g., right direction) were followed by electrical stimulation of the LHb while saccades to the other direction (e.g., left direction) were not. The direction-stimulation contingency was reversed in the next block. We found that the post-saccadic stimulation of the LHb increased the latencies of saccades in subsequent trials. Notably, the increase of the latency occurred gradually as the saccade was repeatedly followed by the stimulation, suggesting that the effect of the post-saccadic stimulation was accumulated across trials. LHb stimulation starting before saccades, on the other hand, had no effect on saccade latency. Together with previous studies showing LHb activation by reward omission and aversive stimuli, the present stimulation experiment suggests that LHb activity contributes to learning to suppress actions which lead to unpleasant events.