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R Romo

Publications and source records attributed to R Romo.

At least 37 records · Page 2Linked to original sources

Representation of moving tactile stimuli in the somatic sensory cortex of awake monkeys.

1. We have studied how neurons of primary somatic sensory (SI) cortex encode the direction and speed of moving tactile stimuli delivered to the glabrous skin of the contralateral hand. 2. From a total of 178 neurons recorded in SI cortex of 3 awake Macaca mulatta monkeys, 103 were selected for quantitative analysis. Forty-six neurons had slowly adapting (SA) responses, 43 quickly adapting (QA), and 14 mixed SA-QA properties. All possessed cutaneous receptive fields on the distal segments of digits 2, 3, or 4. Receptive fields were scanned with a metal probe (2 mm diam, hemispheric tip) in four different directions (0, 90, 180, and 270 degrees), over a fixed traverse distance of 6 mm, at a variety of speeds (4-100 mm/s), and with a static normal force of 20 g by means of a computer-controlled tactile stimulator. 3. Most neurons gave statistically significant differences in mean impulse rate during the moving stimuli (Wilcoxon, P < 0.01), in at least one of the four directions, compared with the control (nonstimulus) period. The Kruskal-Wallis test (P < 0.01) and the direction index (DI > 35%) determined that about one-half of SI cortical neurons showed significant differences in mean impulse rates associated with the direction of the stimuli at the speeds of 23, 50, and 100 mm/s, and about one-third at 4 mm/s. 4. We determined how the temporal covariance of the neural activity was associated with the parameters of the moving stimuli by calculating the coefficients of the Karhunen-Loève (KL) transform for each set of stimulus responses. Decomposition of the neural activity into principal components indicated that approximately 85% of the impulse train variance during the stimulus responses was contained in the 1st 10 coefficients of the KL transform for the speeds of 23, 50, and 100 mm/s, and approximately 75% at the speed of 4 mm/s. The line spectra calculated from the coefficients of the KL transform showed that the variance contained in the impulse trains in about one-half of the neurons is related to the stimuli. 5. We investigated how the temporal covariance of the neuronal activity was correlated with the direction of the stimulus, by fitting the first coefficient of the KL transform to a weighting function model. This analysis showed that the first coefficient of the KL transform varied as an orderly function of the direction of the moving stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

A tactile stimulator for studying motion processing in the somatic sensory system of primates.

A tactile stimulator was built for studying motion processing in the somatic sensory system of primates. This stimulator is used for assessing the responses of neurons of the somatic sensory system to stimuli moving in any traverse distance (range: 2-20 mm), with a variety of velocities (range: 4-120 mm/s), forces (range: 0-60 gf), and in any scanning direction. The stimulator is highly automated and can be used in combined psychophysical and neurophysiological studies in humans and in behaving monkeys.

Animals↗

Representation of tactile signals in primate supplementary motor area.

1. We have studied the neuronal activity in the supplementary motor area (SMA) of two monkeys who categorized the speed of moving tactile stimuli delivered to the glabrous skin of the hand ipsilateral to the site of cortical recording and contralateral to the responding arm. 2. A large number of SMA neurons responded to the stimuli of all speeds (176 of 522) but only when those stimuli controlled behavior. 3. A second class of SMA neurons responded differentially in the categorization task (35 during the stimuli and 51 during the reaction time period) and predicted its outcome. 4. To dissociate the interrupt target switches presses from the tactile categorization responses, sixteen neurons, which responded to the stimuli in all speeds, and 11 neurons, which discharged differentially, were tested in a visual control task. None of these two classes of neurons responded in this situation. 5. It is concluded that the SMA ipsilateral to sensory input and contralateral to the responding arm is involved in the sensory decision process in this somesthetic categorization task.

Afferent Pathways↗

Role of primate basal ganglia and frontal cortex in the internal generation of movements. I. Preparatory activity in the anterior striatum.

The purpose of these studies was to investigate neuronal activity in the basal ganglia and frontal cortex in relation to the internal generation of goal-directed movements. Monkeys performed goal-directed arm movements at a self-chosen moment in the absence of phasic stimuli providing external temporal reference. They were rewarded with a small morsel of food for each movement, although automatic or repetitive behavior was not reinforced. For reasons of comparison, animals were also trained in a delayed go no-go task in which visual cues instructed them to perform or refrain from an arm movement reaction to a subsequent trigger stimulus. This report describes the activity of neurons in the head of the caudate nucleus and rostral putamen preceding self-initiated arm movements and compares it with instruction-induced preparatory activity preceding movements in the delay task. A total of 497 caudate and 354 putamen neurons were tested in the delay task. Two types of preparatory activity were observed: (1) transient responses to the instruction cue, and (2) sustained activity preceding the trigger stimulus or movement onset. Transient responses were found in 48 caudate and 50 putamen neurons, occurring twice as often in movement ('go') as compared to no-movement ('no-go') trials, but rarely in both. These responses may code the information contained in the instruction relative to the forthcoming behavioral reaction. Sustained activity began after instruction onset and lasted until the trigger stimulus or the arm movement occurred, this being for periods of 2-7 s, 12-35 s, or up to 80 s, depending on the task requirements. This activity was seen in 47 caudate and 45 putamen neurons, was largely confined to go trials, and was unrelated to the preparation of saccadic eye movements. In some cases, this activity began as direct responses to the instruction stimulus, but in the majority of cases developed more gradually before the movement. Thus, both transient and sustained activations appear to be related to the preparation of movements. A total of 390 caudate and 293 putamen neurons were tested during self-initiated movements. Activity preceding earliest movement-related muscle activity was found in 32 caudate and 42 putamen neurons. This premovement activity began 0.5-5.0 s before movement onset (median 1160 ms), increased slowly, reached its peak close to movement onset, and subsided rapidly thereafter. It was unrelated to the preparation of saccadic eye movements. Comparisons between the two tasks were made on 53 neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Role of primate basal ganglia and frontal cortex in the internal generation of movements. II. Movement-related activity in the anterior striatum.

In order to more comprehensively assess the role of the basal ganglia in the internal generation of movements, we studied the activity of neurons in the head of the caudate and in the rostral putamen in relation to the execution of movements. Monkeys performed self-initiated and stimulus-triggered arm reaching movements in separate blocks of trials. With stimulus-triggered movements, 217 striatal neurons increased their activity after the trigger stimulus (127 in caudate, 90 in putamen). Of these, 68 neurons showed time-locked responses to the trigger stimulus, with a median latency of 60 ms, that were independent of visual or auditory stimulus modalities. Three quarters of responses were conditional on a movement being performed. These responses may participate in neuronal processes through which the reception of a stimulus is translated into the execution of a behavioral reaction. Further, 44 neurons increased their activity before the earliest muscle activity without being clearly time-locked to the stimulus (148-324 ms before movement onset), 55 neurons were activated later before the movement, and 50 neurons were activated after movement onset. With self-initiated movements, 106 striatal neurons showed movement-related activity beginning up to 460 ms before movement onset (52 in caudate, 54 in putamen). Comparisons between the two types of movement were made on 53 neurons with premovement activity beginning more than 500 ms before self-initiated movements. Only one fifth of them also showed movement-related activity with stimulus-triggered movements, including trigger responses. Comparisons among 39 neurons with movement-related activity during self-initiated arm movements showed that about half of them also showed movement-related activity with stimulus-triggered movements. These data demonstrate a considerably segregated population of striatal neurons engaged in the internal generation of movements, whereas processes underlying the execution of movements appear to involve overlapping neuronal populations.

Animals↗

Role of primate basal ganglia and frontal cortex in the internal generation of movements. III. Neuronal activity in the supplementary motor area.

This study is a part of a project investigating neuronal activity in the basal ganglia and frontal cortex and describes externally and internally induced preparatory activity in the supplementary motor area (SMA), which forms a closed neuronal loop with the striatum. Monkeys made self-initiated arm reaching movements toward a constant target in the absence of phasic external stimuli. In separate blocks of trials, animals performed in a delayed go no-go task in which an instruction cue prepared for subsequent movement or no-movement to a trigger stimulus. A total of 328 neurons were tested in the delay task. Of these, 91 responded transiently to the instruction light with a median latency of 262 ms. Three quarters of these responses were restricted to the instruction preparing for arm movement, as opposed to withholding it, and thus may be involved in movement preparation processes. Sustained activation during the instruction-trigger interval was found for 67 neurons and occurred nearly exclusively in movement trials. Activation usually increased gradually after the cue and ended abruptly upon movement onset and thus could be related to the setting and maintenance of processes underlying the preparation of movement. Time-locked responses to the trigger stimulus were found in 38 neurons and were usually restricted to movement trials (median latency 80 ms). Activity time-locked to movement execution occurred in 67 neurons, beginning up to 252 ms before movement onset. A total of 266 neurons were tested with self-initiated arm movements. Of these, 43 showed premovement activity beginning 610-3030 ms before movement onset (median 1430 ms). The activity increased slowly and reached its peak at 370 ms before movement onset. It ended before movement onset or continued until the arm began to move or reached the target. This activity appears to reflect neuronal processes related to the internal generation of movements. Two thirds of activations preceding self-initiated movements occurred in neurons not activated before externally instructed movements, suggesting a selectivity for the internal generation process. Activity related to the execution of self-initiated movements occurred in 67 neurons: it began during and up to 420 ms before movement onset and was usually not associated with premovement activity. Most of these neurons were also activated with stimulus-triggered movements, suggesting a lack of selectivity for the execution of self-initiated movements. In comparison with the striatum, more SMA neurons showed preparatory activity preceding externally instructed movements (transient 27% vs 16%, sustained 20% vs 12%) and self-initiated movements (16% vs 11%).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Dopamine neurons of the monkey midbrain: contingencies of responses to active touch during self-initiated arm movements.

1. Previous studies have shown that midbrain dopamine (DA) neurons in monkeys respond to external stimuli that are used to initiate behavioral reactions. In the present study, we investigated to what extent changes in neuronal activity would occur when behavioral acts are generated internally or whether they would depend solely on external stimuli. 2. Monkeys performed self-initiated arm movements from a resting key into a covered, food-containing box at a self-chosen moment and without external preparatory or triggering signals. In a second task, the arm movement was triggered by rapid opening of the door of the food box. This stimulus was either audible and visible or only audible to the animal. Impulses of DA neurons were recorded with movable microelectrodes from the pars compacta of substantia nigra (area A9) and areas A8 and A10 and were discriminated from those of other neurons by their long duration (1.5-5.0 ms) and low spontaneous frequency (0.5-8.5 imp/s). 3. The activity of 12% of 104 DA neurons increased slowly and moderately up to 1,500 ms before the onset of individual self-initiated arm movements. Median increases amounted to 91% over background discharge rate. A further 16% of DA neurons were activated together with the onset of muscle activity and during the movement. 4. During self-initiated movements, a nonhabituating, phasic burst of impulses occurred when the monkey's hand touched a morsel of food inside the box. This response was seen in 84% of 154 neurons on the contralateral side, with median onset latency of 65 ms and duration of 160 ms. A comparable percentage of neurons responded to ipsilateral touch with similar latency and duration. 5. The touch response during self-initiated movements was absent, both on the contra- and ipsilateral sides, when the animal's hand touched the bare wire normally holding the food, when touching nonfood objects, or during tactile exploration of the empty interior of the food box. Thus responses appeared to be related to the appetitive properties of the object being touched rather than the object itself. 6. In the task employing stimulus-triggered movements, 77% of 86 DA neurons discharged a burst of impulses in response to door opening but entirely failed to respond to the touch of food in the box. The response to door opening in this task was similar to the touch response during self-initiated movements in the same neurons in terms of latency, duration, and magnitude.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Dopamine neurons of the monkey midbrain: contingencies of responses to stimuli eliciting immediate behavioral reactions.

1. This study investigates the behavioral conditions in which dopamine (DA) neurons of substantia nigra and adjoining areas A8 and A10 respond with impulses to visual and auditory trigger stimuli eliciting immediate arm- and eye-movement reactions. 2. In a formal task, the rapid opening of the door of a small, food-containing box located at eye level ahead of the animal served as visible and audible trigger stimulus. Most DA neurons on the contralateral side responded to this stimulus with a short burst of impulses with median onset latency of 50 ms and duration of 90 ms (75% of 164 neurons). Similar responses were seen in a comparable fraction of DA neurons during ipsilateral task performance, suggesting that responses were not specific for the limb being used. 3. When the sensory components of the door opening stimulus were separated, DA neurons typically responded in a similar manner to the moving visual stimulus of the opening door, the low-intensity sliding noise of the opening door, and the 1-kHz sound of 90-92 dB intensity emitted from a distant source at the onset of door opening. Responses to each component alone were lower in magnitude than to all three together. 4. In a variation of the task, a neighboring, identical food box opened in random alternation with the other box but without permitting animals to reach out (asymmetric, direct-reaction go/no-go task). With each sensory component, DA neurons typically responded both to opening of go and no-go boxes. Responses were enhanced when stimuli elicited limb movements in go trials. 5. Monkeys reacted to door opening with target-directed saccadic eye movements in the majority of both go and no-go trials. Neuronal responses were equally present during the occasional absence of eye movements. Thus responses were not specific for the initiation of individual arm or eye movements. 6. Neuronal responses were absent when the same stimuli occurred outside of the behavioral task with target-direct arm and eye movements lacking. This shows that responses were not of purely sensory nature but were related to the capacity of the stimulus for eliciting behavioral reactions. 7. In a variation of the go/no-go task, an instruction light illuminated 2-3 s before door opening prepared the animal to perform the reaching movement on door opening or to refrain from moving (asymmetric, instruction-dependent go/no-go task).(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗

Frequency discrimination in the sense of flutter: psychophysical measurements correlated with postcentral events in behaving monkeys.

The capacities of humans and monkeys to discriminate between the frequencies of mechanical sinusoids delivered to the glabrous skin of the hand have been measured in psychophysical experiments. The 2 primates have similar capacities; they make discriminations with Weber fractions that change little over the frequency range from 20 to 200 Hz. The discriminatory capacities are similar whether stimuli are received passively or acquired actively. Combined experiments have been made in monkeys in which the electrical signs of the activity of quickly adapting (QA) and slowly adapting (SA) neurons of postcentral areas 3b and 1 were recorded, both in the working state as the animal made discriminations and in the irrelevant state in which the stimuli did not guide behavior. The neuronal responses were analyzed in terms of discharge rates, periodicities in the neuronal discharges, and harmonic contents. It was shown that discriminatory capacity depends upon the period lengths in the sets of periodically entrained activity evoked by stimuli readily discriminated, and not upon the small differences in rates of discharge evoked by those stimuli. The periodicities were shown by harmonic analysis to be sharply limited to stimulus frequencies. Low-frequency stimuli evoke periodicities at the second and third harmonics in some neurons, in addition to strongly periodic signals at the fundamental frequency of the stimuli. Their presence does not appear to interfere with frequency discrimination. Neuronal responses recorded in the stimulus-irrelevant state were not distinguishable from those recorded as monkeys made discriminations. The responses of SA neurons, recorded under similar conditions, resembled those of QA neurons in almost every feature, but reasons are given for concluding that the SA system plays no role in frequency discrimination in the sense of flutter.

Adult↗

Saccadic reaction times, eye-arm coordination and spontaneous eye movements in normal and MPTP-treated monkeys.

The oculomotor performance of monkeys was investigated before and after destruction of nigrostriatal dopamine neurons by MPTP (1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine). Stimulus-triggered saccades and their relationships to arm movements were measured in a reaction time task. Spontaneous eye movements were recorded while monkeys sat in a primate chair and looked around the laboratory without performing any task. In the reaction time task, saccades and arm movements were commonly triggered by the rapid, visible and audible opening of a small food-containing box which was located at a constant position in front of the animal at eye level. Median saccadic latencies ranged from 68 to 118 ms in intact animals. Saccades were followed by onset of electromyographic (EMG) activity in the extensor digitorum communis and the biceps brachii, the prime mover muscles for the following arm reaching movement. Latencies of stimulus-triggered saccades showed an absence of linear relationship to EMG or arm movement reaction time in intact animals (correlation coefficients of 0.15-0.56). This suggests that eye and arm movements were initiated independently from each other in this experimental situation. Treatment with MPTP resulted in 98-99.5% loss of striatal dopamine in both monkeys. This induced a 29-93% increase in saccadic latency in the reaction time task. The sequential occurrence of saccade, EMG activity and arm movement in each trial was preserved, although intervals between onset of saccades and onsets of EMGs and arm movements were prolonged by 53-173% and 33-679% respectively. Onsets of individual saccades remained uncorrelated with onsets of EMG activity or arm movement. Spontaneous eye movements were strongly reduced in frequency and amplitude after MPTP. Administration of the dopamine precursor L-Dopa increased spontaneous eye movements for less than two hours. The severe deficits in stimulus-triggered and spontaneous saccadic eye movements are oculomotor components of hypokinesia arising after MPTP-induced lesions of the nigrostriatal dopamine system in primates. The data are further evidence for a role of midbrain dopamine neurons in behavioral responsiveness and spontaneous activity.

Animals↗

Protection against 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine-induced parkinsonism by the catecholamine uptake inhibitor nomifensine: behavioral analysis in monkeys with partial striatal dopamine depletions.

The neurotoxic effects of 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine on dopamine neurons in monkeys were found to be reduced when the catecholamine uptake inhibitor nomifensine was administered during several weeks after 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine. The obtained protection was partial, leading to 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine-induced changes in dopamine levels to 8, 16, 52 and 59% of control values in the caudate nucleus and to 10, 16, 101 and 99% in the putamen of four animals, respectively. At the same doses, 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine alone is known to deplete striatal dopamine levels to 0.5-7% of control values. Extra-nigrostriatal monoamine neurons were generally well protected by nomifensine. Neurological examinations revealed modest hypokinesia for a maximum of 10 days after 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine in the two more severely affected animals. Reaction times of arm and eye movements were measured in a formal task in two of the monkeys having a moderate and a more important depletion of striatal dopamine, respectively. Only moderate impairments were seen during the initial 2 weeks after 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine in both animals. All parameters recovered to control levels thereafter. At 3.5 and 5.5 months after 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine, task performance was significantly better than control. The speed of arm movement remained largely unaffected during all periods of experimentation. Spontaneous eye movements were reduced in frequency and amplitude during the initial 1-2 weeks after 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine, and recovered completely thereafter. These data suggest a substantial reduction of neurotoxicity of 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine by inhibition of catecholamine uptake. Particularly striking was the absence of major and permanent impairments in behavioral tests in which monkeys treated with 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine alone were severely impaired. These results may warrant the development of new catecholamine uptake inhibitors for protecting nigrostriatal dopamine neurons against potential environmental toxins.

1-Methyl-4-phenyl-1,2,3,6-tetrahydropyridine↗

Somatosensory input to dopamine neurones of the monkey midbrain: responses to pain pinch under anaesthesia and to active touch in behavioural context.

The somatosensory responses of single dopamine (DA) neurones were recorded in the pars compacta of substantia nigra and in neighbouring DA cell groups of four Macaca fascicularis monkeys. These neurones were electrophysiologically discriminated against other cells by their polyphasic, relatively long impulses (2.0-5.0 ms) occurring at low rates (mostly 1.0-5.0/s), by antidromic activation from caudate or putamen, and by reduction of impulse rate following subcutaneous injection of apomorphine (0.05-0.15 mg/kg). Of 140 DA neurones recorded in two monkeys under barbiturate anaesthesia, 51% showed reductions and 17% increases in impulse rate during intense noxious pinch stimulation. Neurones responded non-somatotopically to stimulation of the hand, foot, face, dorsum and tail on both sides of the body. Innocuous, even intense, surface or deep somatosensory stimuli were ineffective. Systemic injection of the DA receptor antagonist haloperidol (0.33-0.5 mg/kg) strongly reduced the pinch responses. Of 154 DA neurones recorded in two monkeys during self-initiated arm movements, 84% showed phasic activations with latencies of 65 ms when the monkey's hand touched a food morsel inside the target box. Responses were absent when touching other objects. Touch responses to food did not occur when the reaching movement into the same food box was performed in reaction to an external trigger stimulus. In conclusion, DA neurones were activated in specific behavioural contexts by somatosensory stimuli of low intensities while responding unconditionally to noxious input.

Action Potentials↗

Deficits in reaction times and movement times as correlates of hypokinesia in monkeys with MPTP-induced striatal dopamine depletion.

1. We quantitatively assessed deficits in the initiation and execution of arm movements occurring after destruction of nigrostriatal dopamine neurons by systemic administration of MPTP (1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine) (Sigma). Three monkeys performed a reaction time task in which they reached toward a single and constant target for food reward. 2. After administration of MPTP, all three monkeys showed hypokinesia necessitating dopamine precursor or receptor agonist treatment. The partial recovery of one animal from initial akinesia after 19 days permitted discontinuation of dopaminergic drug therapy, although marked hypokinesia remained present. The two other animals displayed additional, intermittent phases of rigidity and activation tremor and needed continuous dopaminergic drug therapy for most of the postlesion period. 3. Administration of MPTP significantly prolonged EMG reaction time in prime mover muscles and arm movement reaction time by 47-225% and 18-129%, respectively, on the six sides of the three animals, compared with control measurements before the lesion. EMG and arm movement reaction time increased over consecutive trials in most sessions comprising 110-130 movements, the first 20 movements showing almost normal values. The delay time between onsets of EMG and arm movement showed unsystematic changes. These deficits in movement initiation were observed both with and without dopamine precursor therapy. They lasted during the whole testing period of several months. 4. Linear correlations between arm movement onset and EMG onset in the two prime mover muscles, the extensor digitorum communis and the biceps, showed coefficients of mostly 0.7-0.9, both before and after MPTP. These data suggest that the temporal relationship between onsets of arm movement and EMG were not substantially affected by MPTP. 5. Arm movement time was divided into two phases. The duration of movement between the resting key and the target, a small food-containing box located ahead of the animal, was denoted as reaching movement time. The following hand manipulation inside the food box was measured as box movement time. After MPTP, both measures were significantly prolonged by 10-103% and 12-251%, respectively, on the six sides of the three monkeys. These deficits in movement execution were observed both with and without dopaminergic drug therapy and during the whole testing period. 6. Task performance after MPTP treatment was studied in one monkey in the absence of dopaminergic drug therapy. EMG and arm movement reaction times recovered partially over several weeks, while the prolongations in reaching and box movement times remained unchanged.(ABSTRACT TRUNCATED AT 400 WORDS)

1-Methyl-4-phenyl-1,2,3,6-tetrahydropyridine↗