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Control of single-joint movements in deafferented patients: evidence for amplitude coding rather than position control.

Two deafferented patients and several control subjects participated in a series of experiments to investigate how accurate single-joint movements are programmed, spatially calibrated, and updated in the absence of proprioceptive information. The deafferented patients suffered from a permanent and severe loss of large sensory myelinated fibers below the neck. Subjects performed, with and without vision, sequences of forearm supinations and pronations with two temporal delays between each movement (0 s and 8 s). Overall, the lack of proprioception did not yield any significant decrease in movement accuracy when vision was available. Without vision, the absence of proprioceptive afferents yielded (1) significantly larger spatial errors, (2) amplitude errors similar to those of control subjects, and (3) a significant drift when an 8-s delay was introduced between two successive movements. Subjects also performed, without vision, a 20 degrees supination followed by a 20 degrees pronation that brought back the wrist to the starting position. On some trials, the supination was blocked unexpectedly by way of a magnetic brake. When the supination was blocked, subjects were already on the second target and no pronation was required when the brake was released. The deafferented patients, unaware of the procedure, always produced a 20 degrees pronation. These data confirm that deafferented patients were not coding a final position. It rather suggests that they coded an amplitude and translated the spatial distance between the two targets in a corresponding force pulse. Overall, the results highlight the powerful and key role of proprioceptive afferents for calibrating the spatial motor frame of reference.

Adult↗

Ocular exploration of space as a function of neck proprioceptive and vestibular input--observations in normal subjects and patients with spatial neglect after parietal lesions.

We recently argued that the specific compensation of spatial neglect by manipulating neck proprioceptive and vestibular input is due to a central "correction" of the disturbed neural transformation process converting the afferent input coordinates from the peripheral sensory organs into a central representation of egocentric space. Both types of stimulation were proposed to induce a reorientation of the deviated or distorted egocentric spatial reference frame. The aim of the present study was to observe this process of reorientation under a condition in which no visual stimulus can attract the subject's attention and thus influence exploration behaviour from outside. We recorded eye movements of normal subjects and of three patients with spatial neglect after right parietal lesions while searching for a non-existent target in complete darkness. It was assumed that the area of the outer space that subjects spontaneously explore under this condition is a direct function of the subject's representation of egocentric space. Ocular space exploration was biased and confined almost entirely to the right side of the midsagittal plane in patients with neglect. This spatial distribution of exploratory eye movements changed remarkably with left-sided neck muscle vibration as well as with left-sided vestibular stimulation using ice water calorics. The spatial area of exploration was significantly enlarged to the contralesional side and the exploration maximum shifted in the same direction. Whereas with both types of stimulation space exploration of patients with neglect was similar to that of normal subjects when not being stimulated, neck proprioceptive and vestibular stimulation in normal subjects induced a quasi neglect-like exploration pattern, i.e. a bias to one side of the objective midsagittal plane. If ocular space exploration was, however, related to the subjectively perceived position of the midsagittal plane in space, eye movements were symmetrically distributed and carried out to both sides of subjective "straight ahead" in all experimental conditions, in normal subjects as well as in patients with neglect. The present results support the above hypothesis and indicate that neck proprioceptive as well as vestibular input directly contribute to the computation of the subject's central representation of egocentric space used for localizing body orientation and for guiding motor behaviour in space.

Adult↗

Static spatial effects in motor cortex and area 5: quantitative relations in a two-dimensional space.

We describe the relations between active maintenance of the hand at various positions in a two-dimensional space and the frequency of single cell discharge in motor cortex (n = 185) and area 5 (n = 128) of the rhesus monkey. The steady-state discharge rate of 124/185 (67%) motor cortical and 105/128 (82%) area 5 cells varied with the position in which the hand was held in space ("static spatial effect"). The higher prevalence of this effect in area 5 was statistically significant. In both structures, static effects were observed at similar frequencies for cells that possessed as well as for those that lacked passive driving from the limb. The results obtained by a quantitative analysis were similar for neurons of the two cortical areas studied. It was found that of the neurons with a static effect, the steady-state discharge rate of 78/124 (63%) motor cortical and 63/105 (60%) area 5 cells was a linear function of the position of the hand across the two-dimensional space, so that the neuronal "response surface" was adequately described by a plane (R2 greater than or equal to 0.7, p less than 0.05, F-test in analysis of variance). The preferred orientations of these response planes differed for different cells. These results indicate that individual cells in these areas do not relate uniquely a particular position of the hand in space. Instead, they seem to encode spatial gradients at certain orientations.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Goal-directed arm movements in absence of visual guidance: evidence for amplitude rather than position control.

The control of pointing arm movements in the absence of visual guidance was investigated in unpracticed human subjects. The right arm grasped a lever which restricted the movement of the right index fingertip to a horizontal arc, centered between the axes of eye rotation. A horizontal panel directly above the arm prevented visual feedback of the movement. Visual stimuli were presented in discrete positions just above panel and fingertip. A flag provided visual feedback on fingertip position before each pointing movement (Exp. A and B), or before a movement sequence (Exp. C). When subjects pointed from straight ahead to eccentric stimulus positions (Exp. A), systematic and variable pointing errors were observed; both kinds of errors increased with stimulus eccentricity. When subjects pointed from 30 deg left to stimuli located further right (Exp. B), errors increased with stimulus position to the right. Taken together, these findings suggest that pointing accuracy depends not primarily on stimulus position, but rather on required movement amplitude. When subjects performed sequences of unidirectional movements (Exp. C), systematic and variable errors increased within the sequence. A quantitative analysis revealed that this increase can be best described as an accumulation of successive pointing errors. We conclude that both findings, error increase with amplitude, and accumulation of successive errors, when considered together strongly support the hypothesis that amplitude, rather than final position, is the controlled variable of the investigated movements.

Arm↗

Effects of posterior parietal lesions (area 7) on VOR in monkeys.

Unilateral ablations of area 7 were performed in three adult monkeys. Vestibulo-ocular reflex (VOR) was tested in the dark by sinusoidal rotations at different frequencies. Following lesion of area 7, spontaneous nystagmus was observed in the dark, with the fast phase directed toward the lesioned side. The same lesion induced a strong VOR asymmetry due to a gain decrease when the animal was rotated toward the side contralateral to the lesion and an increase when rotated toward the opposite side. These VOR deficits were transient: spontaneous nystagmus was no longer present after the first post-operative week whereas the VOR asymmetry lasted for 2 to 4 weeks after the lesion. It is concluded that area 7 might be involved in an ipsilateral control of the slow component of VOR. These results support the idea that posterior parietal cortex plays a role in body reference stabilization.

Animals↗

Spatial attention and eye movements.

We previously showed that when attention is allocated to the right or left of the fixation point, saccades directed to targets located above or below the fixation point deviate contralateral to the attention locus. In the present study, we examined how general this phenomenon is and whether the amount of saccade deviation depends on the location of attention with respect to that of the saccade target. Three experiments were carried out. In experiment 1 the location of the imperative stimulus was uncued. Its presentation exogenously directed attention to its location. In experiment 2 the location of the imperative stimulus was cued by a central cognitive cue. In this experiment attention was endogenously directed to the imperative stimulus location before its presentation (expectancy paradigm). In experiment 3 all stimulus boxes contained a possible imperative stimulus at the display presentation. A central cue, presented subsequently, indicated which of them had to be used for the saccade. In this experiment attention was endogenously directed to the imperative stimulus, but after its presentation (no-expectancy paradigm). The results showed that, regardless of how attention was directed to the imperative stimulus, the vertical saccades deviated contralateral to the attention location. The deviation was larger when attention was in the upper field and the saccade was directed upward ("same hemifield" condition) than when attention was in the upper field and the saccade was directed downward ("opposite hemifield" condition). The same relationship between the "same hemifield" condition and "opposite hemifield" condition was found when attention was in the lower field. Saccadic reaction times (SRTs) were shortest in experiment 2 and longest in experiment 3. In experiment 2, SRTs of the "same hemifield" condition were significantly longer than those of the "opposite hemifield" condition. Taken altogether, these results strongly support the notion that attention allocation in space leads to an activation of oculomotor circuits, in spite of eye immobility. The possible mechanisms responsible for saccade deviations and for greater saccade deviations when attention is in the same hemifield as the programmed ocular saccade are discussed.

Analysis of Variance↗

Moving one's finger to a visually specified position: target orientation influences the finger's path.

It has previously been shown that, when subjects are instructed to move their finger slowly from one point to another the finger follows a path that deviates systematically from a straight line connecting the two points. The deviation depends on the angle between this fictive line and a line connecting the subject's finger with his body. In the present study, we examined whether the deviation also depends on the target's orientation. In two experiments, subjects were instructed to move a finger slowly towards five targets. We recorded the finger's movements. In one experiment, the targets were aligned. In the other, they were oriented radially around the starting point. Otherwise, conditions were the same. The difference in target orientation influenced the finger's path. Most importantly, when the targets were oriented radially around the starting point, the finger's path was straight. We conclude that pointing is more than moving the finger to a specified position.

Fingers↗

The nucleus accumbens in monkeys (Macaca fascicularis). III. Reversal learning.

The nucleus accumbens (NA), which receives inputs from limbic structures and projects to the motor system, may be important for the association of reinforcement with action. There are projections to the NA from the amygdala and hippocampus. Discrimination and reversal learning tasks which are known to be disrupted by lesions to these areas in monkeys were given to monkeys with lesions of the NA. Twelve monkeys (Macaca fascicularis) were used in the present study. Six of these received ibotenic acid lesions which resulted in considerable cell loss in the NA; the remaining six acted as controls. The first group of six monkeys were taught a visual discrimination task pre-operatively. Post-operatively, these monkeys were tested on visual and spatial discrimination and reversal tasks. A second group of six monkeys were tested on a motor reversal task. The results indicate that ibotenic acid lesions of the NA transiently impair spatial but not visual reversal learning in monkeys. The NA lesions did not impair a monkey's ability to perform visual or spatial discriminations, or the ability to perform the motor learning or motor reversal tasks. Our results suggest that bilateral lesions of the NA in monkeys do not disrupt the ability to discriminate basic properties of reward-related stimuli or the formation of visual stimulus-reward associations. In addition, our results argue against theories which suggest that the NA is important for behavioural switching or general behavioural flexibility. We conclude that the NA may play a more specific role in the association of temporal and spatial cues with movement and reward.

Animals↗

Effects of deprivation of vision and vibrissae on goal-directed locomotion in cats.

The locomotor behaviour of adult cats blinded soon after birth was compared to that of control cats by measuring running time in a maze. The effects of vibrissae deprivation in trained blind and control cats and of visual deprivation in control cats was studied. The performance of the control cats decreased in darkness to the level reached by the blind cats. Deprivation of vibrissae affected the performance of the control cats only in darkness and had no effect on the blind cats. The role played by various sensory systems in locomotion during development and in adulthood is discussed.

Animals↗

A functional analysis of the collicular output pathways: a dissociation of deficits following lesions of the dorsal tegmental decussation and the ipsilateral collicular efferent bundle in the Mongolian gerbil.

Mongolian gerbils received either lesions of the superior colliculus, small lesions of the uncrossed efferents of the superior colliculus, knifecuts of the dorsal tegmental decussation, or sham operations. The animals were subsequently tested for avoidance of overhead visual threat, orientation and locomotion toward small targets, and negotiation of a large barrier in order to reach a small target. Animals with collicular lesions showed no responses to overhead threat and had severe deficits in orienting to small targets. Animals with lesions of the uncrossed tectal efferents showed diminished responses to overhead threat but had no deficits in orienting to targets. Animals with lesions of the dorsal tegmental decussation showed only slight reductions in responses to overhead threats but clear impairments in the orientation tasks. The impairments in orientation, however, were less severe than those seen in collicular animals. Animals in all groups were able to negotiate barriers efficiently. These results suggest that separate output pathways of the superior colliculus mediate different types of visuomotor behaviours. The results further suggest that visual orientation to small targets does not depend completely on output through the predorsal bundle, but must also involve other collicular outputs.

Animals↗

Influences of lesions of parietal cortex on visual spatial attention in humans.

Several brain areas have been identified with attention, because damage to these regions leads to neglect and extinction. We have tested elements of visual attentional processing in patients with parietal, frontal, or temporal lesions and compared their responses to control subjects. Normal humans respond faster in a reaction time task when the spatial location of a target is correctly predicted by an antecedent stimulus (valid cue) than when the location is incorrectly predicted (invalid cue). The cue is hypothesized to shift attention towards its location and thereby facilitate or impede response latencies. The reaction times of individuals with damage to the parietal lobe are somewhat slowed for targets ipsilateral or contralateral to the side of the lesion if the targets are preceded by valid cues. These same patients are extremely slow in responding to targets in the visual field contralateral to the lesion when the cue has just appeared in the unaffected (ipsilateral) visual field. In addition, these individuals are especially slow in responding to targets in either visual field when the lights are preceded by weak, diffuse illumination of the entire visual field. Patients with lesions of the frontal lobe have very slow reaction times in general and, as is the case for patients with lesions of the temporal lobe, are slow in all conditions for targets in the field contralateral to the lesion. These patterns are probably not associated with attentional defects. For patients with parietal lesions, these studies demonstrate a further deficit in a cued reaction-time task suggesting abnormal visual attention. Since different sites of brain damage yield different patterns of responses, tests such as these could be of analytic and diagnostic value.

Adolescent↗

Effects of fornix transection and cingulate cortical ablation on spatial memory in rhesus monkeys.

This study, together with a parallel study in rats (Markowska et al. 1988), attempted to relate the effects of hippocampal-system damage on similar tasks in both rats and monkeys. Not only were monkeys given a task (Experiment 1) which was of the sort usually used with rats, but in the companion study rats were given tasks (Experiment 2) like those usually used with monkeys. Experiment 1 examined the performance of rhesus monkeys with hippocampal-system damage on a spatial working memory task. Monkeys were trained preoperatively on delayed nonmatching-to-sample in a T-maze, placed into groups matched for their preoperative learning scores, and then received one of three treatments: 1) transection of the fornix; 2) ablation of the cingulate cortex; or 3) a sham operation. Monkeys with fornix transection were severely and significantly impaired, but monkeys with cingulate cortical ablations were not significantly impaired, relative to the controls. The results demonstrate that monkeys with fornix transection are severely impaired on a spatial working memory task requiring locomotion and, taken together with earlier work, suggest that the effect of fornix transection in both rodents and nonhuman primates is at least qualitatively similar (see Markowska et al. 1988). Experiment 2 assessed the role of the fornix and cingulate cortex in three conditional tasks in which the monkeys were provided with various spatial cues to indicate which one of two objects was rewarded. Both experimental groups were unimpaired, relative to the control group, on all three tasks, indicating that fornix transection does not produce a general impairment in place learning.

Animals↗

A comparative analysis of the role of fornix and cingulate cortex in memory: rats.

In order to compare the role of the fornix (FX) and cingulate cortex (CC) in memory, rats were trained in a series of discriminations using procedures that were the same as those used for monkeys (Murray et al. 1986, 1988). A spatial delayed nonmatching-to-sample (DNMS) discrimination tested recent memory for spatial location in a T maze using interrun intervals (IRI) that varied from 5 s to 15 min. FX and CC lesions produced a substantial impairment in the performance of this task during postoperative testing. Three conditional discriminations (CD) followed. In each one, the rat was presented with two objects, only one of which was correct. The nature of the conditional stimuli changed in each discrimination: the place of the maze in the room; the direction that the rat moved to approach the objects; the side (left or right) to which the rat turned. Control rats learned all three types of conditional discriminations. FX and CC lesions did not impair choice accuracy. In a subsequent repetition of the spatial DNMS procedure, FX and CC lesions again produced a substantial impairment, indicating that the lack of an impairment in the three CDs was not due to recovery of function. These data indicate that the hippocampal system and its connections through the fornix are importantly involved in spatial working memory in both rats and monkeys, and that the CDs do not require this type of memory. The results are discussed in the context of different theories of the brain mechanisms involved in memory.

Animals↗

Place memory and scene memory: effects of fornix transection in the monkey.

Five experiments examined the effects of fornix transection upon some spatial and visual learning tasks in monkeys (Macaca fascicularis). For each trial of each task, the monkey was brought to a test tray and allowed to choose between 2 objects on the tray. In different tasks, different cues were provided by the experimenter to guide the monkey's choices. In total 5 different tasks were run (Experiments 1 to 5) and the results showed that the effects of fornix transection varied markedly between tasks: the animals with fornix transection were severely impaired in experiments 1, 3 and 5 but learned normally in experiments 2 and 4. It is concluded that the results cannot be explained by the simple hypothesis of a deficit in place learning, since some forms of place learning are unimpaired by fornix transection. A better general hypothesis is that the memory disrupted by fornix transection is like a snapshot memory, which stores the spatial arrangement of items in a witnessed scene.

Animals↗

Multivalued stimulus-response relation in isolated elasmobranch utricles.

The relation between a maintained spatial orientation and the corresponding fully adapted discharge rate was multivalued in all the afferents tonically sensitive to maintained spatial orientation observed in isolated utricles of Rhinobates productus. The spread of rate values was of the order of changes produced by natural tilts. The occurrence of multivaluedness in isolated receptors indicated that peripheral issues are sufficient. Two factors contributed: firstly, the side from which the orientation had been reached (i.e. "hysteresis"): higher adapted rates occurred when the preceding orientation was characterized by lower rates and when the corresponding transition caused acceleration; secondly, "spontaneous" rate variations, some of which resembled markedly, and interacted with, the effects of tilts. It was not possible to identify the basic mechanisms underlying these factors. The multivaluedness in the coding of maintained position, because of its constancy and magnitude, cannot be ignored. It, as well as the sensitivity to fast transients, must be taken into account in utricular models, in evaluations of information transmission, and in psychophysical explorations.

Action Potentials↗

Learning spatial sequences in unilateral neglect.

Brain-damaged patients with unilateral spatial neglect ignore aspects of the world located on the side opposite their lesion. In the present study we examined the performance of unilateral neglect patients (UN) on an SRT task in which a hybrid repeating sequence (21313) was used. We analyzed the patients' performance for each location separately as a function of the target's location in the trial preceding the response. The UN patients were severely limited in their learning of the sequence when compared to normal controls. In particular, they appeared to learn unique associations (21 and 13) but not ambiguous ones (31 and 32). We discuss two possible explanations for this phenomenon. The first is that UN patients show a deficit similar to that of normal subjects in dual task situations. The second is that the learning deficit is unique to spatial processing impairments of UN patients and is not directly related to research with normal population. We outline future research that may distinguish between these two explanations.

Attention↗