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E P Gardner

Publications and source records attributed to E P Gardner.

At least 19 recordsLinked to original sources
PubMed2001

Comparison of neuronal firing rates in somatosensory and posterior parietal cortex during prehension.

To evaluate their functional roles during prehension, single-unit recordings were made in the hand area of primary somatosensory areas 3b, 1 and 2 (S-I) and posterior parietal areas 5 and 7 (PPC) of the same animal. Response profiles of mean firing rate during performance of a multistage reach, grasp, and lift task were analyzed to determine the period(s) of peak firing and to measure statistically significant rises or falls in rate compared with baseline. We used the peak firing stage(s) to subdivide the population into classes tuned to single actions or two successive stages, or into multiaction groups that had sustained facilitation (BT) or inhibition (GI) during hand-object interactions. Four times as many neurons fired at peak rates during acquisition stages (approach, contact, grasp) than upon release, and their firing rates were higher. Grasping evoked the strongest responses, as grasp-tuned neurons had the highest peak rates in the population; BT, contact-grasp, and grasp-lift cells also fired maximally in the grasp stage. Grasping also coincided with maximal inhibition of GI cells, as well as of neurons tuned to approach or relaxation of grasp. Holding evoked the lowest mean rates, and had the fewest tuned cells. S-I and PPC showed significant differences in behaviors evoking peak firing as well as facilitation and inhibition; these correlated with input modalities in each area. Hand contact with the object and positioning of the fingers for grasp was the most strongly represented behavior in anterior S-I, where 61% received tactile inputs from glabrous skin. Nearly 60% were facilitated at contact, 38% fired at peak rates, and 10% were inhibited; release of grasp evoked peak firing in only 5% of 3b-1 neurons. In posterior S-I, where proportions of tactile and deep inputs were similar, positioning and grasping elicited peak responses in 38% and 31%, respectively; 80% were facilitated or inhibited during grasping. During lift and hold, inhibition rose to 43%, while excitation declined under 10%. PPC had the highest proportions firing at peak rates during hand preshaping before contact (28%) and had the most facilitated responses (38%) in this stage. Only 10% fired at peak rates during grasping. During later manipulatory actions, proportions of facilitated and inhibited responses in PPC were similar to those in posterior S-I. The data support models in which PPC plans hand movements during prehension rather than guiding their execution. Sensory monitoring of hand-object interaction occurs in S-I, where cells sense specific hand behaviors, signal stage completion, enable error correction, and may update grasp programs formulated in PPC. The results are discussed in relation to those obtained from lesion studies in humans.

Animals↗
PubMed2000

Depression of neuronal firing rates in somatosensory and posterior parietal cortex during object acquisition in a prehension task.

Prehension is an object-oriented behavior consisting of four components: reach, grasp, manipulation, and release. To determine how such actions are represented in primary somatosensory (S-I) and posterior parietal cortex (PPC), we used digital video to synchronize spike trains of neurons recorded in Brodmann's areas 3b, 1, 2, 5, and 7 with the hand kinematics as monkeys performed a prehension task. Statistical analyses indicated that one-third of task-modulated neurons showed significantly depressed firing rates during object acquisition and/or manipulation. This population was dominated by neurons innervated by deep receptors that sensed extension movements of the fingers, or by tactile receptors in hairy skin sensing stretch. Grasp-inhibited responses were the most common type. Tonic firing rates of these cells dropped significantly during approach as the hand was preshaped for grasping, or at contact when grasp was initiated, and persisted until hand motion ceased or as the grip relaxed. Maximum suppression of firing occurred at grasp completion. Their lack of specificity for particular hand behaviors formed the inhibitory counterpart of broadly tuned cells that fired prolonged bursts during grasp and manipulatory stages of prehension. The remainder of the task-inhibited population showed biphasic responses. Firing rates were significantly depressed during grasping and manipulation when the hand interacted directly with the object, but were enhanced prior to contact, when the hand was preshaped (approach-tuned), or upon relaxation of grasp and release of the object from the hand (loweror relax-tuned). Grasp-inhibited responses occurred primarily in S-I, whereas biphasic inhibitory activity was recorded mainly in PPC. Suppression of activity within these populations may thereby increase the saliency of excitatory responses to acquisition and manipulation of objects. Reduction of firing during prehension might also signal the flexed postures used to retain objects in the hand, rather than a generalized gating of sensory information. The similarity of responses to active and passive extension movements suggests that the inhibitory responses may provide important postural and motor information about the hand kinematics when performing skilled tasks.

Action Potentials↗
PubMed1999

Facilitation of neuronal activity in somatosensory and posterior parietal cortex during prehension.

In order to study prehension in a reproducible manner, we trained monkeys to perform a task in which rectangular, spherical, and cylindrical objects were grasped, lifted, held, and lowered in response to visual cues. The animal's hand movements were monitored using digital video, together with simultaneously recorded spike trains of neurons in primary somatosensory cortex (S-I) and posterior parietal cortex (PPC). Statistically significant task-related modulation of activity occurred in 78% of neurons tested in the hand area; twice as many cells were facilitated during object acquisition as were depressed. Cortical neurons receiving inputs from tactile receptors in glabrous skin of the fingers and palm, hairy skin of the hand dorsum, or deep receptors in muscles and joints of the hand modulated their firing rates during prehension in consistent and reproducible patterns. Spike trains of individual neurons differed in duration and amplitude of firing, the particular hand behavior(s) monitored, and their sensitivity to the shape of the grasped object. Neurons were classified by statistical analysis into groups whose spike trains were tuned to single task stages, spanned two successive stages, or were multiaction. The classes were not uniformly distributed in specific cytoarchitectonic fields, nor among particular somatosensory modalities. Sequential deformation of parts of the hand as the task progressed was reflected in successive responses of different members of this population. The earliest activity occurred in PPC, where 28% of neurons increased firing prior to hand contact with objects; such neurons may participate in anticipatory motor control programs. Activity shifted rostrally to S-I as the hand contacted the object and manipulated it. The shape of the grasped object had the strongest influence on PPC cells. The results suggest that parietal neurons monitor hand actions during prehension, as well as the physical properties of the grasped object, by shifting activity between populations responsive to hand shaping, grasping, and manipulatory behaviors.

Animals↗
PubMed1998

Digital video: a tool for correlating neuronal firing patterns with hand motor behavior.

This report describes the use of multimedia technology for simultaneous recording of single unit responses in cerebral cortex, and imaging of hand kinematics as monkeys grasp and manipulate objects. These imaging methods allow direct correlation of full-frame, full-field video images with the actual spike trains recorded with microelectrodes. Our implementation of digital video provides high-resolution snapshots of the hand motor behavior every 33.3 ms, and a precise calibration and display of the synchronously recorded electrophysiological activity digitized at rates up to 44.5 kHz on the same platform. These imaging methods permit non-invasive, non-traumatic monitoring of both trained and spontaneous activity in experimental animals, while providing synchronized digitized records of neuronal spike trains. We also describe software instruments that quantify and analyze the digitized spike trains. One instrument employs user-selectable objective criteria for distinguishing spikes from noise, separates individual action potential waveforms by their amplitude and duration, and compiles time stamps for each spike train. A second instrument constructs rasters and histograms of repeated behavioral trials using the timing of the corresponding video frame for alignment. These analyses reveal functional classes of cortical neurons signaling specific stages of prehension.

Animals↗
PubMed1996

Discrimination of simulated texture patterns on the human hand.

1. Textures formed by periodic dot arrays are defined by the dot density, spacing, and angular orientation with respect to the direction of motion. In this report we evaluate the effects of the dot density (intensive cues) and arrangement (spatial cues) on the ability of human subjects to discriminate texture patterns scanned across an OPTACON tactile stimulator that selective stimulates rapidly adapting cutaneous mechanoreceptors. We compared dot arrays arranged on the index finger in specific patterns (horizontal, vertical, diamond, up diagonal, or down diagonal orientation) and spaced 4.8, 7.2, or 9.6 mm apart (high, medium, and low density) with the use of a two-alternative forced-choice protocol. 2. Textures are well discriminated when their elements are tightly spaced along one axis and widely spaced on all other axes. Humans distinguish textures that differ only in orientation with mean accuracy of 75% at low density and 65% at medium density, but discriminate high-density textures poorly (mean accuracy = 48%). Accuracy is related to the angular disparity between patterns, and to similarity of spacing and orientation along major and minor axes of the arrays. Vertical and horizontal patterns are more accurately distinguished than the oblique ones, and diamond arrays are the least well discriminated. Diagonal and diamond textures are often confounded, and the up and down diagonal patterns are confused with each other particularly as the texture density rises. The preference for the vertical and horizontal patterns may relate to an interaction between the orientation axis of the texture and its direction of motion across the skin. 3. Intensive cues provided by the total number of applied stimuli supplement the spatial cues inherent to the pattern orientation, because textures that differ in both spacing and orientation are discriminated better than those that differ only in orientation or spacing. Mean accuracy ranges from 96% for comparisons of high- and low-density textures, which differ in the total number of dots by a factor of 2, to 80% when medium-density patterns are compared with high- or low-density textures. 4. Textures that differ in density but not in orientation are less well discriminated than those of different orientation. For example, 82% of patterns that differ in both density and orientation are distinguished correctly in pairings of low- and medium-density textures, whereas those that differ only in density are discriminated correctly on 45% of trials. Subjects seem to use spatial rather than intensive cues when discriminating patterns of similar density, suggesting that the similarity of form (the spatial arrangement of the closely spaced dots) is more readily apparent than small differences in spacing along the axis of motion. 5. Subjects are most most successful in differentiating texture patterns when they are able to mentally picture the orientation and spacing of the pattern. We found a strong correlation between the subjects' ability to discriminate textures of a given spacing and their ability to identify the specific texture by matching it to the appropriate visual representation. Subjects are able to identify correctly all five orientations at low and medium densities, with mean accuracy of 76%, but recognize only the vertical arrays when high-density patterns are presented. The ability to image the textures is noteworthy, because subjects received no feedback about performance. 6. Spatial imaging of textures appears limited by the diameter of cutaneous receptive fields on the hand. We propose that the structural axis of a regular texture array results from perceptual linkage of adjacent elements along one principal axis by continuous bands of neural activity when their spacing is smaller than the receptive field diameter.(ABSTRACT TRUNCATED)

Adult↗
PubMed1994

Discrimination of the direction of motion on the human hand: a psychophysical study of stimulation parameters.

1. In these experiments we assess the relative importance of the spatial and temporal properties of a moving tactile stimulus in determining the ability of humans to discriminate its direction of motion. Movement along the finger was simulated by applying a series of pulses to adjacent locations on the skin using the tactile array of an OPTACON stimulator. Simulated motion permitted us to vary independently the overall distance moved as well as the spacing, timing, and number of sequential stimuli. Different combinations of spatiotemporal parameters allowed us to further examine the relationship of apparent velocity of motion and sweep duration to behavioral performance. Discrimination accuracy was measured using signal detection techniques to calculate the discrimination parameter d' and PCmax, a bias-free measure of the percent correct identification of the direction of motion. 2. In experiments where the path length was constant, discriminability of the direction of motion increased as the spacing between successive pulses narrowed. Similarly, for a given interpulse spacing, the accuracy of discrimination increased linearly with distance, saturating at perfect performance. These apparent spatial effects on performance actually reflect the total number of stimuli presented to the skin rather than their proximity. Sweeps containing the same number of pulses are equally discriminable regardless of either their spacing or the total distance crossed on the skin. d' values obtained at 1.2-, 2.4-, and 4.8-mm spacings appear indistinguishable when plotted as a function of the total number of pulses in a sweep. 3. Experiments in which both the distance moved and the spacing between pulses was varied randomly confirmed that discrimination accuracy depends on the total number of pulses in a sweep rather than the spatial dimensions of the path traversed. Stimulation of only two points that mark the start and stop locations on the skin appears insufficient to enable subjects to discriminate correctly the direction of motion. Two-point stimulation elicits random performance whether the points lie 1.2 or 4.8 mm apart. Discriminability rises linearly to near-perfect performance when eight or more pulses are delivered sequentially. Extrapolation of the d' and PCmax curves suggests a mean threshold of approximately three points for 75% correct discrimination of the direction of motion across the skin. 4. The relationship of stimulus spacing to discriminability over a fixed path suggests that direction discrimination does not simply involve computation of the location of the start and stop points on the skin or their spatial disparity.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗
PubMed1992

Simulation of motion on the skin. V. Effect of stimulus temporal frequency on the representation of moving bar patterns in primary somatosensory cortex of monkeys.

1. To assess the mechanisms used by cortical neurons to sense motion across the skin, we applied pulsatile stimuli to a series of adjacent positions on the glabrous skin of the hand using a computer-controlled OPTACON stimulator. We describe responses of 129 single neurons in primary somatosensory cortex of alert monkeys to a horizontal bar pattern that was displaced proximally or distally in 1.2-mm steps at 10-, 20-, and 40-ms intervals (100, 50, and 25 Hz, respectively). These frequencies span the range in which apparent motion is transformed perceptually in humans from a smooth uninterrupted sweep into a series of distinct pulses that are resolved as separate events. The experiments are thus designed to decipher the neural correlates distinguishing continuous motion from discrete taps. 2. Cortical receptive fields mapped with moving bar patterns spanned 5-24 rows on the tactile array (16.2 +/- 5.4, mean +/- SD). Over 40% of the fields encompassed 18 or more rows (greater than or equal to 21.6 mm), allowing these neurons to integrate spatial information from an entire image displayed on the OPTACON. Cortical receptive fields are considerably larger than those of mechanoreceptors mapped with the same moving bar patterns (4.2 +/- 2.3 rows, mean +/- SD), reflecting convergent inputs in subcortical and cortical relays. Responses were either relatively uniform across the field or strongest at the initial point of entry, depending on the frequency of stimulation. A sharply defined field center was absent from most of the cells recorded in this study. 3. Temporal frequency of stimulation appears to be a major determinant of cortical firing patterns. Low-frequency stimuli are more effective in activating cortical neurons, producing more spikes per sweep and greater phase-locking to individual stimuli than do high frequencies. The total spike output of cortical neurons is proportional to the pulse interval over the range 10-40 ms, increasing linearly by an average of 5.9 spikes/10-ms increase in pulse period. Peak firing rates and modulation amplitude are also highest when pulses are presented at long intervals, falling significantly as the stimulation frequency rises. The reduction in firing at high pulse rates is apparently due to central mechanisms, as both rapidly adapting and Pacinian corpuscle afferents display nearly constant spike outputs and uniform sensitivity within the field when tested with identical bar patterns. Central networks thus behave as low-pass filters, reducing cortical responses to rapidly applied sequential stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1990

Simulation of motion on the skin. III. Mechanisms used by rapidly adapting cutaneous mechanoreceptors in the primate hand for spatiotemporal resolution and two-point discrimination.

1. The contribution of rapidly adapting (RA) mechanoreceptors to two-point discrimination has been evaluated by examining their ability to resolve the spacing of grating patterns shifted across the skin. The experiments test two different neural coding mechanisms that have been proposed to underlie resolution of spatial detail on the hand: 1) a rate-intensity code in which the spacing of surface features is encoded by the average frequency of firing of individual sensory afferents, and 2) an isomorphic representation of shape in which variations in the firing patterns of individual afferents reflect the spatiotemporal profile of skin indentation. 2. To measure the spatial acuity of RA mechanoreceptors innervating the hands of macaque monkeys, we displayed pairs of horizontal bars spaced 1-13 mm apart on a computer-controlled OPTACON stimulator placed over glabrous skin. Two-point resolution was measured by simultaneously pulsing pairs of rows at rates of 100, 50 and 25 Hz; each pair was shifted in tandem across the hand to simulate lateral motion. Single-fiber recordings were made from physiologically identified RA afferents in anesthetized monkeys. 3. Receptive field diameter appears to be the critical determinant of spatial resolution of gaps between two bars. RAs fire continuously if bar spacing is less than the field diameter but do not summate inputs when both active rows are contained within the field. Response profiles evoked by two bars spaced less than 4.8 mm apart can be predicted from the single-bar profiles, assuming occlusion between overlapping inputs with the strongest member dominating axonal output. Two-thirds of the RAs tested discharge 1 spike/pulse as bar patterns cross the field, yielding a uniform spike train whose frequency reflects stimulus pulse rates but fails to indicate gaps between bars. An additional 17% fire 2 spikes/pulse when the bars contact or straddle the field center, but also fail to differentiate individual stripes spaced less than 3.6 mm apart. 4. Only 17% of RAs represent gaps narrower than the field diameter. These fibers show double-peaked response profiles to bar patterns spaced at least 2.4 mm apart, firing 2 spikes/pulse as first one, and then the second stripe crosses the field center. Timing between peaks corresponds to bar spacing. Responses are reduced in amplitude when adjacent bars straddle the field center, as occlusion between simultaneous inputs prevents summation of inputs from the two stimuli. Fifteen of 16 RAs failed to resolve bars spaced 1.2 mm apart, as double-spike responses were evoked only by the leading stripe.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1990

Simulation of motion on the skin. IV. Responses of Pacinian corpuscle afferents innervating the primate hand to stripe patterns on the OPTACON.

1. To measure spatial acuity of Pacinian corpuscle (PC) afferents in the median and ulnar nerves of macaque monkeys, we displayed horizontal bar patterns spaced 1-13 mm apart on a computer-controlled OPTACON stimulator contacting the hand. Two-point resolution was measured by simultaneously pulsing pairs of rows at rates of 100, 50, and 25 Hz; each pair was shifted in tandem across the skin in 1.2-mm steps to simulate tangential motion at speeds of 30-120 mm/s. Single-fiber responses are reported from eight physiologically identified PC afferents innervating the fingers and palm in anesthetized monkeys. 2. Pacinian afferents differ in their sensitivity to stripe patterns moved across the hand. Bursting PCs fire bursts of two or three spikes/pulse when one of the bars is close to the field center and one spike/pulse when adjacent bars straddle the center. These bursts result in double-peaked response profiles at stripe spacings greater than or equal to 2.4 mm. The passage of individual stripes over the field center is therefore represented by bursts of impulses superimposed on a continuous spike train. Unfortunately, many of these fibers also demonstrate fluctuations in firing that appear unrelated to the stripe pattern and therefore obscure its clear representation. 3. The remainder of the PC population displays uniform-sensitivity responses that resemble those previously reported for rapidly adapting (RA) afferents. They fire one spike/pulse as long as at least one of the bars is contained within the field. They merge individual stripes spaced less than one field diameter apart and show a pause in firing at wider spacing. Spatial resolution of gaps in the stripe pattern is therefore determined by receptive-field diameters, which extend up to 9.6 mm when tested with the OPTACON. 4. PCs display poorer spatial resolution than RAs, because of their larger receptive fields and less regular firing patterns. Only two of eight PCs tested demonstrated a pause in activity representing the gap between bars spaced 4.8 mm apart, whereas 11 of 14 RAs ceased firing briefly between stripes. Resolution of the individual stripes by all of the PCs tested was observed only at bar spacings of 1 cm (8 rows) or more. Spatial resolution of stripes is further impeded by the tendency of PC afferents to summate inputs from stripes spaced less than 2.4 mm apart; this results in response profiles with a single, large-amplitude broad peak.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1989

Simulation of motion on the skin. I. Receptive fields and temporal frequency coding by cutaneous mechanoreceptors of OPTACON pulses delivered to the hand.

1. Tactile discrimination of form requires motion of the hand across the object scanned. To dissociate lateral distortion of the skin from neuronal processing mechanisms involving multiple receptor classes and parallel central networks, we have simulated motion of bar patterns across the fingers and palm by the use of a computer-controlled grid of sequentially activated probes (OPTACON stimulator). Horizontal bar patterns have been swept across the hand at speeds of 30-120 mm/s to quantitatively characterize responses of cutaneous mechanoreceptive afferents recorded in the median and ulnar nerves. 2. Mechanoreceptors with phasic responses to pressure are activated by spatial patterns on the OPTACON, whereas those with tonic pressure responses are not; moving-bar patterns strongly excite both Meissner's afferents [rapidly adapting (RA) mechanoreceptors] and Pacinian corpuscles (PCs) but fail to excite slowly adapting (SA) afferents. OPTACON-type stimulators thus allow selective activation of phasic mechanoreceptor channels with spatially complex stimuli. 3. RA afferents respond in an all-or-none fashion to activation of two to five adjacent rows spanning 1-5 mm on the finger, with nearly identical latencies on all trials; response profiles are remarkable for their regularity and reproducibility. PCs have larger fields (4-13 rows) and stronger but more irregular responses than RAs. 4. Uniform sensitivity throughout the receptive field is a consistent feature of RA responses. Individual mechanoreceptor terminals appear to have equal access to the spike initiation zone and provide the same amplitude input as the fiber discharges 1 spike/pulse at each field location in 75% of the RAs tested. Uniform sensitivity allows each afferent to transmit a repetitive signal of the parameter of interest such as object speed, contour, or texture. 5. One-quarter of RAs fire two spikes to probe indentation and retraction at the field center. Such graded responses are usually observed in only one direction of motion, reflecting a preferred sequence of receptor activation rather than a specific location on the skin. PCs fire bursts of two to four spikes throughout most of their receptive fields; sensitivity is broadly distributed rather than peaked. Thus phasic mechanoreceptors fail to provide a precise signal of stimulus location; localization at the level of individual papillary ridges appears to be signaled by a population mechanism involving unique combinations of RA, SA, and PC afferents.(ABSTRACT TRUNCATED AT 250 WORDS)

Action Potentials↗
PubMed1989

Simulation of motion on the skin. II. Cutaneous mechanoreceptor coding of the width and texture of bar patterns displaced across the OPTACON.

1. These experiments assay the functional significance of receptive-field architecture for information processing. Rapidly adapting (RA) afferents have been previously shown to converge information from clusters of 14-25 Meissner's corpuscles, whereas afferents innervating Pacinian corpuscles (PCs) have only a single, large receptor terminal. We tested two opposing hypotheses of functional architecture: 1) summation models, in which an afferent integrates signals from all of its terminals, showing monotonic increases in activity as a function of contact area, and 2) winner-take-all models, in which the most strongly activated receptor in the cluster dominates axonal output by cancellation of signals from other branches. 2. Bar and stripe patterns have been swept across the finger or palm of the monkey's hand at speeds of 30-120 mm/s with the use of a computer-controlled grid of sequentially activated miniature probes (OPTACON stimulator). The dense packing of OPTACON probes permits placement of up to five groups of stimulators within an individual receptive field, allowing us to activate one or more clusters of Meissner's corpuscles simultaneously and to stimulate the bulbar corpuscle of PC afferents at different orientations through the skin. Integration of information from moving bar patterns has been tested with two protocols. In the variable width protocol, the total number of activated rows in the pattern is varied from one to five, with a constant spacing of 1.2 mm between pulsed rows. In the variable density protocol, the length of skin stimulated is held constant at 5 mm and the spacing of stimuli varied. 3. RA afferents show no evidence of summation of inputs within their receptive fields. Motion of wide bars across the field increases the duration of firing but not the total spikes evoked by each pulse. Responses to the leading edge of wide bars were found to be identical to those evoked by a single-row bar. Simultaneous activation of two to five rows evokes the same or fewer spikes per pulse than the most effective individual row tested alone. When broad-bar patterns are centered over the field, contacting the maximum number of receptors, RAs follow activity in the dominant branch or terminus, suppressing additional inputs. Lack of summation is observed at all pulse frequencies tested (25-100 Hz). 4. Moving bar patterns evoke responses as long as at least one row stimulates the receptive field; broader patterns evoke longer spike trains whose total number of impulses is proportional to bar width.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1989

OAHC opposed.

Explore the source record for details and available documents.

Allied Health Personnel↗
PubMed1988

Somatosensory cortical mechanisms of feature detection in tactile and kinesthetic discrimination.

Neurons in somatosensory cortex of primates process sensory information from the hand by integrating information from large populations of receptors to extract specific features. Tactile neurons in areas 1 and 2 are shown to select features such as contact area, edge orientation, motion across the skin, or direction of movement. Features coded by kinesthetic neurons in areas 3a and 2 relate to joint movement, the joint angle around which the movement occurs, or coordinated postures of the hand and arm. An even higher order cortical cell integrates tactile and kinesthetic information; these "haptic neurons" respond optimally to contact of objects actively grasped in the hand. These global features are coded at the expense of loss of information concerning fine-grained spatial detail.

Animals↗
PubMed1986

Objective classification of motion- and direction-sensitive neurons in primary somatosensory cortex of awake monkeys.

In order to classify movement-sensitive neurons in SI cortex, and to estimate their relative distribution, we have developed a new simple method for controlled motion of textured surfaces across the skin, as well as a set of objective criteria for determining direction selectivity. Moving stimuli were generated using 5 mm thick precision gear wheels, whose teeth formed a grafting. They were mounted on the shafts of low-torque potentiometers (to measure the speed and direction of movement) and rolled manually across the skin using the potentiometer shaft as an axle. As the grafting wheel was advanced, its ridges sequentially contacted a specific set of points on the skin, leaving gaps of defined spacing that were unstimulated. This stimulus was reproducible from trial to trial and produced little distention of the skin. Three objective criteria were used to categorize responses: the ratio of responses to motion in the most and least preferred directions [direction index (DI)], the difference between mean firing rates in the two directions divided by the average standard deviation [index of discriminability (delta'e)], and statistical tests. Neurons were classified as direction sensitive if DI greater than 35, delta's greater than or equal to 1.35 (equivalent to 75% correct discrimination by an unbiased observer), and firing rates in most- and least-preferred directions were significantly different (P less than 0.05). Good agreement was found between the three classification schemes. Recordings were made from 1,020 cortical neurons in the hand and forearm regions of primary somatosensory cortex (areas 3b, 1 and 2) of five macaque monkeys. Tangential motion across the skin was found to be an extremely effective stimulus for SI cortical neurons. Two hundred eighty six of 757 tactile neurons (38%) responded more vigorously to moving stimuli than to pressure or tapping the skin. One hundred twenty-one cells were tested with moving gratings and were classified according to their ability to differentiate movement in longitudinal and transverse directions. Responses to the moving gratings resembled those observed when stroking the skin with brushed, edges, or blunt probes. Three major types of firing patterns were found: motion sensitive, direction sensitive, and orientation sensitive. Motion-sensitive neurons (37%) responded to movement in both longitudinal and transverse directions with only slight difference in firing rates and interval distributions. Responses throughout the field were fairly uniform, and no clear point of maximum sensitivity was apparent. Direction-sensitive neurons (60%) displayed clear preferences for movement in one or more directions.4

Action Potentials↗
PubMed1986

Coding of the spatial period of gratings rolled across the receptive fields of somatosensory cortical neurons in awake monkeys.

In order to measure the texture coding capabilities of motion-, direction-, and orientation-sensitive neurons in SI cortex, we rolled wheels with surface milled gratings across their receptive fields. Gratings of spatial periods 0.8-9.6 mm were presented in pseudorandom order; each was tested 5-20 times in the distal, proximal, radial, and ulnar directions. Thirty eight cortical neurons were studied with three to eight different gratings in order to determine the effect of spatial period on neuronal firing rates. While all 38 cells had their firing rates modulated by motion of the gratings, only 11 neurons were able to distinguish changes in its spatial period. These cells had small receptive fields located on the hand. Most motion-sensitive neurons showed little effect of spatial period on firing rates and had relatively flat frequency response curves. One showed decreased firing to spatial periods over the range 0.8-6.4 mm; three others increased their firing rates over the range 0.8-3.2 mm, followed by a decline in activity to larger spatial periods. Direction- and orientation-sensitive neurons showed only minor changes in firing rates as a function of spatial period. Sixteen cells showed flat frequency response functions, three showed increased firing rates, and four decreased firing rates as spatial period of the grating increased. Direction and orientation preferences were maintained over the range 0.8-9.6 mm for all 23 neurons tested. Although four cells showed a drop in direction index (DI) as the spatial period was increased, none showed a loss of direction sensitivity, as DI was greater than 35 for all gratings tested. Two neurons showed increased firing to motion in the last-preferred direction and two others decreased firing in the best direction. The remaining 19 neurons showed parallel effects of texture in all directions. Some motion-sensitive neurons showed weak direction preferences when tested with fine gratings; these preferences disappeared with coarser gratings, due to increased responsiveness to motion in the least-preferred direction. These data demonstrate that movement-sensitive neurons do not require continuous trajectories across the skin but instead sequential activation of points aligned in a specific path. Cortical neurons appear capable of integrating information from points separated by up to 9 mm, as long as they are presented in the appropriate temporal sequence. Firing rates of direction- and orientation-sensitive neurons are more profoundly modified by changes in the direction of motion across the skin, and the temporal order of stimulation, than by alterations in the spatial characteristics of the moving stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗
PubMed1985

Differential sensitivity to airpuffs on human hairy and glabrous skin.

To compare the relative sensitivities of glabrous and hairy skin, we measured reaction times (RTs) and detectability (d') of airpuffs delivered to the hairy dorsum and glabrous thenar eminence of the hand of six human subjects. In contrast to previous studies with mechanical contact stimuli, airpuffs applied to hairy skin were detected with equal or greater fidelity than airpuffs tested on glabrous skin. Mean RTs to three simultaneously applied airpuffs were significantly shorter (p less than .005) on hairy skin in five of six subjects, and in 74% of paired sessions; no significant difference in mean RTs was observed in 16% of the sessions. The superiority of hairy skin was less evident, however, when single airpuffs were tested, as significantly shorter responses were observed on only 45% of the paired sessions, and nearly identical responses on 38% of the sessions. Detectability of airpuffs (d'), which is independent of the value of RTs, was identical on hairy and glabrous skin at high airpuff intensities (1,600 dyn), and superior (n = 4) or equal (n = 2) on hairy skin with low airpuff intensities (800 dyn). Spatial summation was more pronounced on hairy than on glabrous skin. Three simultaneously presented airpuffs produced significantly shorter RTs than one airpuff in 85% of the paired sessions on hairy skin, but on only half of the sessions on glabrous skin. The spatial distribution of stimulus force was less important on hairy skin, as three low-intensity airpuffs produced the same or shorter RTs than one high-intensity airpuff. By contrast, on glabrous skin, detectability was significantly better when force was concentrated at a single point (1 X 1,600 dyn) than when diffused over a wide skin area (3 X 800 dyn). The enhanced sensitivity of hairy skin to airpuffs appears partially attributable to hair motion in the airstream. After hair removal by chemical depilation, detectability of airpuffs was reduced on hairy skin to a level equal to or below that on glabrous skin. Spatial summation on the depilated skin corresponded to that observed on the intact hairy skin, indicating that depilation did not abolish intensity discrimination, but rather lowered the overall sensitivity of hairy skin. These results show that hair follicle units form a very sensitive detection mechanism on hairy skin of the human hand, similar to that provided by Meissner's and Pacinian afferents in glabrous skin. These findings with airpuffs provide the first example of a tactile stimulus that is less effective for mechanoreceptors in glabrous skin than in hairy skin.(ABSTRACT TRUNCATED AT 400 WORDS)

Adult↗
PubMed1984

Somatosensory evoked potentials (SEPs) and cortical single unit responses elicited by mechanical tactile stimuli in awake monkeys.

The origins of surface recorded evoked potentials have been investigated by combining recordings of single unit responses and somatosensory evoked potentials (SEPs) from the postcentral gyrus of 4 alert macaque monkeys. Responses were elicited by mechanical tactile stimuli (airpuffs) which selectively activate rapidly adapting cutaneous mechanoreceptors, and permit patterned stimulation of a restricted area of skin. Epidurally recorded SEPs consisted of an early positive complex, beginning 8-10 msec after airpuff onset, with two prominent positive peaks (P15 and P25), succeeded by a large negative potential (N43) lasting 30 msec, and a late slow positivity (P70). SEPs, while consistent in wave form, varied slightly between monkeys. The amplitude of the early positive complex was enhanced by increasing the number of stimulated points, or by placing the airpuffs in the receptive fields of cortical neurons located beneath the SEP recording electrode. SEP amplitude was depressed when preceded 20-40 msec earlier by a conditioning stimulus to the same skin area. Single unit responses in areas 3b and 1 of primary somatosensory (SI) cortex consisted of a burst of impulses, beginning 11-12 msec after the airpuff onset, and lasting another 15-20 msec. Peak unitary activity occurred at 12-15 msec, corresponding to the P15 wave in the SEP. No peak in SI unit responses occurred in conjunction with the P25 wave. Although SI neurons fired at lower rates during P25, the lack of any peak in SI unit responses suggests that activity in other cortical areas, such as SII cortex, contributes to this wave. Most unit activity in SI cortex ceased by the onset of N43, and was replaced by a period of profound response depression, in which unit responses to additional tactile stimuli were reduced. We propose that the N43 wave reflects IPSPs in cortical neurons previously depolarized and excited by the airpuff stimulus. Late positive potentials (P70) in the SEP had no apparent counterpart in SI unit activity, suggesting generation at other cortical loci.

Animals↗
PubMed1984

Two epizootics of lymphocytic choriomeningitis virus occurring in laboratory mice despite intensive monitoring programs.

Two epizootics of lymphocytic choriomeningitis virus in mice occurred within two months in one research facility consisting of several widely separated rooms. These outbreaks developed despite intensive institutional monitoring policies designed to prevent introduction and spread of lymphocytic choriomeningitis virus. Evidence derived from serological and virological assays and interviews with the concerned investigators suggested that a single transplantable tumor carried in mice may have been responsible for spread of the virus. However, the tumor was not contaminated with lymphocytic choriomeningitis virus at the time of its introduction into the mouse facility. The origin of the virus responsible for the outbreaks was not definitively established although data supported an hypothesis that the virus was introduced into the research facility by a wild or feral mouse. Virus spread from infected mice to humans did not occur, as measured by serological tests. However, a large and valuable animal facility was depopulated for safety reasons. Absorption of sera with lymphocytic choriomeningitis virus antigen proved a necessary and reliable method for confirming specificity of lymphocytic choriomeningitis virus fluorescence-positive reactions.

Animals↗