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I Salimi

Publications and source records attributed to I Salimi.

7 recordsLinked to original sources

Specificity of internal representations underlying grasping.

The present study examines anticipatory control of fingertip forces during grasping based on the center of mass (CM) of a manipulated object. Subjects lifted an object using a precision grip while the fingertip forces and the angle about the vertical axis (roll) were measured. The object's CM could be shifted to the left or right of the object's center parallel to the grip axis without changing it's visual appearance. Subjects performed 20 lifts with the CM in the center, left, and right side of the object, respectively. Subjects were instructed to lift the object while preventing it from tilting. Within three to five lifts, subjects were able to asymmetrically partition the load force development before lift-off such that it was higher in the digit opposing the CM. This anticipatory load force partitioning prevented the object from rolling sideways at lift-off. To determine whether the internal representation underlying the anticipatory control is specific to the effectors used to form it, subjects performed five lifts with the right hand with the CM on one side. Following these lifts, they rotated the object 180 degrees around the vertical axis and performed one lift with the same hand or they translated the object to the left side of the body (with or without rotating it) and performed one lift with the left hand. Despite subjects' explicit knowledge of the new weight distribution, they were unable to appropriately scale the load forces at each digit, resulting in a subsequent large roll of the object. The findings suggest that within a few lifts subjects achieve a stable internal representation which accounts for the object's CM and is used to scale the fingertip forces in advance. They also suggest that this representation, which is used for anticipatory control of fingertip forces, is specific to the effectors used to form it. We propose that multiple internal representations may be used during the anticipatory control of grasping.

Adult↗

Neuronal activity in somatosensory cortex of monkeys using a precision grip. I. Receptive fields and discharge patterns.

Three adolescent Macaca fascicularis monkeys weighing between 3.5 and 4 kg were trained to use a precision grip to grasp a metal tab mounted on a low friction vertical track and to lift and hold it in a 12- to 25-mm position window for 1 s. The surface texture of the metal tab in contact with the fingers and the weight of the object could be varied. The activity of 386 single cells with cutaneous receptive fields contacting the metal tab were recorded in Brodmann's areas 3b, 1, 2, 5, and 7 of the somatosensory cortex. In this first of a series of papers, we describe three types of discharge pattern, the receptive-field properties, and the anatomic distribution of the neurons. The majority of the receptive fields were cutaneous and covered less than one digit, and a chi2 test did not reveal any significant differences in the Brodmann's areas representing the thumb and index finger. Two broad categories of discharge pattern cells were identified. The first category, dynamic cells, showed a brief increase in activity beginning near grip onset, which quickly subsided despite continued pressure applied to the receptive field. Some of the dynamic neurons responded to both skin indentation and release. The second category, static cells, had higher activity during the stationary holding phase of the task. These static neurons demonstrated varying degrees of sensitivity to rates of pressure change on the skin. The percentage of dynamic versus static cells was about equal for areas 3b, 2, 5, and 7. Only area 1 had a higher proportion of dynamic cells (76%). A third category was identified that contained cells with significant pregrip activity and included cortical cells with both dynamic or static discharge patterns. Cells in this category showed activity increases before movement in the absence of receptive-field stimulation, suggesting that, in addition to peripheral cutaneous input, these cells also receive strong excitation from movement-related regions of the brain.

Action Potentials↗

Neuronal activity in somatosensory cortex of monkeys using a precision grip. II. Responses To object texture and weights.

Three monkeys were trained to lift and hold a test object within a 12- to 25-mm position window for 1 s. The activity of single neurons was recorded during performance of the task in which both the weight and surface texture of the object were systematically varied. Whenever possible, each cell was tested with three weights (15, 65, and 115 g) and three textures (smooth metal, fine 200 grit sandpaper, and rough 60 grit sandpaper). Of 386 cells recorded in 3 monkeys, 45 cells had cutaneous receptive fields on the index or thumb or part of the thenar eminence and were held long enough to be tested in all 9 combinations of texture and weight. Recordings were made for the entire anterior-posterior extent of the thumb and index finger areas in somatosensory cortex including area 7b. However, the statistical analysis required a selection of only those cells for which nine complete recording conditions were available limiting the sample to cells in areas 2, 5, and 7b. Significant differences in the grip force accompanied 98% of the changes in texture and 78% of the changes in weight. Increasing the object weight also increased the force tangential to the skin surface as measured by the load or lifting force. The peak discharge during lifting was judged to be the most sensitive index of cell activity and was analyzed with a two-way analysis of variance (ANOVA). In addition, peak cell discharge was normalized to allow comparisons among different combinations of texture and weight as well as comparisons among different neurons. Overall, the peak firing frequency of 87% of the cells was significantly modulated by changes in object texture, but changes in object weight affected the peak activity of only 58% of the cells. Almost all (17/18, 94%) of the static cells were influenced by the object texture, and 81% of the dynamic cells that were active only briefly at grip and lift onset were modulated by texture. For some cells, surface texture had a significant effect on neuronal discharge that was independent of the object weight. In contrast, weight-related responses were never simple main effects of the weight alone and appeared instead as significant interactions between texture and weight. Four neurons either increased or decreased activity in a graded fashion with surface structure (roughness) regardless of the object weight (P < 0.05). Ten other neurons showed increases or decreases in response to one or two textures, which might represent either a graded response or a tuning preference for a specific texture. The firing frequency of the majority (31/45) of neurons reflected an interaction of both texture and weight. The cells with texture-related but weight-independent activities were thought to encode surface characteristics that are largely independent of the grip and lifting forces used to manipulate the object. Such constancies could be used to construct internal representations or mental models for planning and controlling object manipulation.

Afferent Pathways↗

Neuronal activity in somatosensory cortex of monkeys using a precision grip. III. Responses to altered friction perturbations.

The purpose of this investigation was to examine the activity changes in single units of the somatosensory cortex in response to lubricating and adhesive coatings applied to a hand-held object. Three monkeys were trained to grasp an object between the thumb and index fingers and to lift and hold it stationary within a narrow position window for 1 s before release. Grip forces normal to the skin surface, load forces tangential to the skin surface, and the displacement of the object were measured on each trial. Adhesive (rosin) and lubricant (petroleum jelly) coatings were applied to the smooth metal surface of the object to alter the friction against the skin. In addition, neuronal activity evoked by force pulse-perturbations generating shear forces and slip on the skin were compared with the patterns of activity elicited by grasping and lifting the coated surfaces. Following changes in surface coatings, both monkeys modulated the rate at which grip forces normal to the skin surface and load forces tangential to the skin surface were applied during the lifting phase of the task. As a result, the ratio of the rates of change of the two forces was proportionately scaled to the surface coating properties with the more slippery surfaces, having higher ratios. This precise control of normal and tangential forces enabled the monkeys to generate adequate grip forces and prevent slip of the object. From a total of 386 single neurons recorded in the hand area of the somatosensory cortex, 92 were tested with at least 1 coating. Cell discharge changed significantly with changes in surface coating in 62 (67%) of these cells. Of these coating-related cells, 51 were tested with both an adhesive and lubricating coating, and 45 showed significant differences in activity between the untreated metal surface and either the lubricant or the adhesive coating. These cells were divided into three main groups on the basis of their response patterns. In the first group (group A), the peak discharge increased significantly when the grasped surface was covered with lubricant. These cells appeared to be selectively sensitive to slip of the object on the skin. The second group (group B) was less activated by the adhesive surface compared with either the untreated metal or the lubricated surface, and they responded mainly to variations in the force normal to the skin surface. These cells provide useful feedback for the control of grip force. The third group (group C) responded to both slips and to changes in forces tangential to the skin. Most of these cells responded with a biphasic pattern reflecting the bidirectional changes in load force as the object was first accelerated and then decelerated. One hundred sixty-eight of the 386 isolated neurons were tested with brief perturbations during the task. Of these, 147 (88%) responded to the perturbation with a significant change in activity. In most of the cells, the response to the perturbation was shorter than 100 ms with a mean latency of 44.1 +/- 16.3 (SD) ms. For each of the cell groups, the activity patterns triggered by the perturbations were consistent with the activity patterns generated during the grasping and lifting of the coated object.

Animals↗

The effects of peripheral deafferentation on spontaneously bursting neurons in the somatosensory cortex of waking cats.

Single neurons (n = 356) were studied in the forelimb representation of awake, quietly resting cats. Thirty-five spontaneously bursting neurons in a sample of 206 cells recorded before forelimb deafferentation were compared to 39 spontaneously bursting neurons in a sample of 127 neurons studied 1-3 weeks after deafferentation. The probability of encountering bursting neurons increased significantly following deafferentation from 17% to 31% of the sample (P < 0.005). The same 5 classes of bursting cells were observed after deafferentation but there were significant changes in the duration of interspike intervals in some classes, in the probability of observing certain classes, and in the proportion of spikes found in bursts. The probability of encountering class III cells, a class thought to consist primarily of non-inactivating pyramidal burst neurons, nearly doubled and the average interspike interval length within the burst increased from 1.9 to 3.0 ms. The burst structure in the other classes did not change but they were found less frequently. These other classes may include inhibitory interneurons which receive less excitatory drive after deafferentation and therefore provide less inhibition to class III cells. The differential behavior of the different classes of bursting cells may be one reason why the overall level of spontaneous activity does not change after deafferentation and it suggests that there are homeostatic mechanisms in primary somatosensory cortex that maintain a certain level of neural activity.

Afferent Pathways↗

Neuronal activity in normal and deafferented forelimb somatosensory cortex of the awake cat.

Three hundred and seventy-three neurons were recorded from the forelimb representation in the primary somatosensory cortex of unanesthetized, quietly resting adult cats. Of these, 177 were studied from 2 days to 3 weeks after transection of the radial, median and ulnar nerves. Following deafferentation the proportion of cells without receptive fields increased from 24 to 82%, however, the average rate of spontaneous activity did not change nor did the probability of encountering a neuron with a receptive field as a function of depth. Receptive field sizes increased dramatically following deafferentation and the response changed from a reliable short-latency, brisk discharge to one that did not occur on every stimulus. After deafferentation the edges of the receptive field often could not be defined accurately. Spontaneous activity in 31% (n = 47) of the neurons from deprived cortex could be modulated by manipulations of the body but these changes were sufficiently slow and ill-defined that they were not classified as a receptive field. In some cases, manipulation of the body gradually reduced the discharge rate. This slow decline in activity was different from the abrupt inhibition of spontaneous activity elicited by somatic stimuli in another class of cells (n = 18). In other cases the manipulation produced a gradual increase in the discharge rate. After deafferentation antidromically identified corticothalamic and pyramidal tract neurons did not display behaviors different from their counterparts in normal cortex. However, the mean latency for synaptic activation from the ventroposterior thalamus increased from 2.7 ms to 4.6 ms. The lost forelimb receptive fields were rarely replaced by inputs from adjacent body parts over the two-week duration of this study. Most responses to somatic stimuli obtained from cortical neurons in the deafferented cortex were clearly abnormal.

Afferent Pathways↗

Reevaluation of area 3b in the cat based on architectonic and electrophysiological studies: regional variability with functional and anatomical consistencies.

Most anatomical and electrophysiological studies of the cat primary somatosensory cortex rely on Hassler and Muhs-Clement's (J. Hirnforsch. 6:377-420, 1964) cyto- and myeloarchitectonic description distinguishing area 3a from area 3b; however, discrepancies in the delineation of these areas in published studies suggest that many workers have found it difficult to apply those criteria systematically. We examined the cytoarchitecture of area 3b in Nissl stained sagittal sections from which electrophysiological data had been obtained prior to sacrifice. Rostrocaudal rows of electrode penetrations placed at different mediolateral positions in the gyrus located regions responsive to stimulation of either cutaneous or deep structures. Small electrolytic lesions allowed these data to be related to the cytoarchitecture. A systematic study throughout the trunk and limb representations found cutaneous responses in cortical regions characterized by a thick and cell-dense granular layer IV, however these same regions had a variable population of medium-sized and/or large pyramidal cells in layer V. Pyramidal cells were practically absent from the forelimb representation, but were present to varying degrees in the trunk and hindlimb representations. Moreover, the relative thickness and cell-density in layer IV were greater in the forelimb than in the hindlimb representations. Deep responses were found in cortex characterized by a thinner layer IV. Since the characteristics of layer V in area 3a were variable, it was less useful for identification of the border between areas 3a and 3b. Clear changes in the intensity and laminar distribution of acetylcholinesterase staining occurred between areas 3a and 3b, making this a useful adjunct to the Nissl stain.

Acetylcholinesterase↗