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K O Johnson

Publications and source records attributed to K O Johnson.

At least 19 recordsLinked to original sources
PubMed2001

Neural coding mechanisms underlying perceived roughness of finely textured surfaces.

Combined psychophysical and neurophysiological studies have shown that the perceived roughness of surfaces with element spacings of >1 mm is based on spatial variation in the firing rates of slowly adapting type 1 (SA1) afferents (mean absolute difference in firing rates between SA1 afferents with receptive fields separated by approximately 2 mm). The question addressed here is whether this mechanism accounts for the perceived roughness of surfaces with element spacings of <1 mm. Twenty triangular and trapezoidal gratings plus a smooth surface were used as stimulus patterns [spatial periods, 0.1-2.0 mm; groove widths (GWs), 0.1-2.0 mm; and ridge widths (RWs), 0-1.0 mm]. In the human psychophysical studies, we found that the following equation described the mean roughness magnitude estimates of the subjects accurately (0.99 correlation): 0.2 + 1.6GW - 0.5RW - 0.25GW(2). In the neurophysiological studies, these surfaces were scanned across the receptive fields of SA1, rapidly adapting, and Pacinian (PC) afferents, innervating the glabrous skin of anesthetized macaque monkeys. SA1 spatial variation was highly correlated (0.97) with human roughness judgments. There was no consistent relationship between PC responses and roughness judgments; PC afferents responded strongly and almost equally to all of the patterns. Spatial variation in SA1 firing rates is the only neural code that accounts for the perceived roughness of surfaces with finely and coarsely spaced elements. When surface elements are widely spaced, the spatial variation in firing rates is determined primarily by the surface pattern; when the elements are finely spaced, the variation in firing rates between SA1 afferents is determined by stochastic variation in spike rates.

Action Potentials↗
PubMed2001

Differences in spatial acuity between digits.

The spatial acuity of the index, middle, and ring distal finger pads of eight right-handed men, aged 22 to 57 years, was measured by using gratings and raised letters. Acuity declined significantly from the index to the middle finger and from the middle to the ring finger. There were no significant differences between homologous fingers of the two hands. Letter recognition and grating orientation threshold measures were highly correlated.

Adult↗
PubMed2001

The roles and functions of cutaneous mechanoreceptors.

Combined psychophysical and neurophysiological research has resulted in a relatively complete picture of the neural mechanisms of tactile perception. The results support the idea that each of the four mechanoreceptive afferent systems innervating the hand serves a distinctly different perceptual function, and that tactile perception can be understood as the sum of these functions. Furthermore, the receptors in each of those systems seem to be specialized for their assigned perceptual function.

Animals↗
PubMed2000

Attention modulates synchronized neuronal firing in primate somatosensory cortex.

A potentially powerful information processing strategy in the brain is to take advantage of the temporal structure of neuronal spike trains. An increase in synchrony within the neural representation of an object or location increases the efficacy of that neural representation at the next synaptic stage in the brain; thus, increasing synchrony is a candidate for the neural correlate of attentional selection. We investigated the synchronous firing of pairs of neurons in the secondary somatosensory cortex (SII) of three monkeys trained to switch attention between a visual task and a tactile discrimination task. We found that most neuron pairs in SII cortex fired synchronously and, furthermore, that the degree of synchrony was affected by the monkey's attentional state. In the monkey performing the most difficult task, 35% of neuron pairs that fired synchronously changed their degree of synchrony when the monkey switched attention between the tactile and visual tasks. Synchrony increased in 80% and decreased in 20% of neuron pairs affected by attention.

Action Potentials↗
PubMed2000

Spatial and temporal structure of receptive fields in primate somatosensory area 3b: effects of stimulus scanning direction and orientation.

This is the third in a series of studies of the neural representation of tactile spatial form in somatosensory cortical area 3b of the alert monkey. We previously studied the spatial structure of >350 fingerpad receptive fields (RFs) with random-dot patterns scanned in one direction () and at varying velocities (). Those studies showed that area 3b RFs have a wide range of spatial structures that are virtually unaffected by changes in scanning velocity. In this study, 62 area 3b neurons were studied with three to eight scanning directions (58 with four or more directions). The data from all three studies are described accurately by an RF model with three components: (1) a single, central excitatory region of short duration, (2) one or more inhibitory regions, also of short duration, that are adjacent to and nearly synchronous with the excitation, and (3) a region of inhibition that overlaps the excitation partially or totally and is temporally delayed with respect to the first two components. The mean correlation between the observed RFs and the RFs predicted by this three-component model was 0.81. The three-component RFs also predicted orientation sensitivity and preferred orientation to a scanned bar accurately. The orientation sensitivity was determined most strongly by the intensity of the coincident RF inhibition in relation to the excitation. Both orientation sensitivity and this ratio were stronger in the supragranular and infragranular layers than in layer IV.

Action Potentials↗
PubMed2000

Neural coding.

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Animals↗
PubMed2000

Tactile functions of mechanoreceptive afferents innervating the hand.

Four types of mechanoreceptive afferents innervate the glabrous skin of the hand. Evidence from more than three decades of combined psychophysical and neurophysiological research supports the idea that each afferent type serves a distinctly different sensory function and that these functions explain most of tactual perceptual function. The available evidence supports the following hypotheses: (1) The slowly adapting type 1 system provides the information on which form and texture perception are based. (2) The cutaneous rapidly adapting system provides information about minute skin motion and, thereby, plays a critical role in grip control. (3) The Pacinian system is responsible for the detection and perception of distant events by vibrations transmitted through objects, probes, and tools held in the hand. (4) The slowly adapting type 2 system provides information for the perception of hand conformation and for the perception of forces acting on the hand. The authors review the evidence on which these hypotheses are based. They also review the role of proprioceptive afferents in the perception of hand conformation because they appear to play a significant role along with cutaneous afferents.

Afferent Pathways↗
PubMed2000

The afferent neural response to electrotactile stimuli: preliminary results.

We recorded action potentials from three RA fibers innervating primate fingerpad, while applying electrotactile (electrocutaneous) stimulation. Negative pulses required 1.3-1.5 times more current than positive ones for entrainment. The strength-duration time constant was approximately 151 micros. Suprathreshold sinusoidal vibration synchronized to 30-Hz electrotactile pulses changed the electrotactile entrainment current slightly, indicating a possible electrical-mechanical transduction interaction.

Action Potentials↗
PubMed1999

Velocity invariance of receptive field structure in somatosensory cortical area 3b of the alert monkey.

This is the second in a series of studies of the neural representation of tactile spatial form in cortical area 3b of the alert monkey. We previously studied the spatial structure of 330 area 3b neuronal receptive fields (RFs) on the fingerpad with random dot patterns scanned at one velocity (40 mm/sec; ). Here, we analyze the temporal structure of 84 neuronal RFs by studying their spatial structure at three scanning velocities (20, 40, and 80 mm/sec). As in the previous study, most RFs contained a single, central, excitatory region and one or more surrounding or flanking inhibitory regions. The mean time delay between skin stimulation and its excitatory effect was 15.5 msec. Except for differences in mean rate, each neuron's response and the spatial structure of its RF were essentially unaffected by scanning velocity. This is the expected outcome when excitatory and inhibitory effects are brief and synchronous. However, that interpretation is consistent neither with the reported timing of excitation and inhibition in somatosensory cortex nor with the third study in this series, which investigates the effect of scanning direction and shows that one component of inhibition lags behind excitation. We reconcile these observations by showing that overlapping (in-field) inhibition delayed relative to excitation can produce RF spatial structure that is unaffected by changes in scanning velocity. Regardless of the mechanisms, the velocity invariance of area 3b RF structure is consistent with the velocity invariance of tactile spatial perception (e.g., roughness estimation and form recognition).

Adaptation, Physiological↗
PubMed1999

Detection of vibration transmitted through an object grasped in the hand.

A tool or probe often functions as an extension of the hand, transmitting vibrations to the hand to produce a percept of the object contacting the tool or probe. This paper reports the psychophysical results of a combined psychophysical and neurophysiological study of the perception of vibration transmitted through a cylinder grasped in the hand. In the first part of the psychophysical study, 19 subjects grasped a cylinder, 32 mm diam, with an embedded motor that caused vibration parallel to the axis of the cylinder. The relationship between threshold and frequency was the traditional U-shaped function with a minimum between 150 and 200 Hz. Except a study by Békésy in which subjects grasped a rod that vibrated parallel to the skin surface, thresholds above 20 Hz were lower and the slopes were steeper than any reported previously. Thresholds were <0.01 microm in some subjects. Data from both the psychophysical and the neurophysiological studies suggest that detection performance at frequencies >20 Hz was based on activity in Pacinian afferents. The extreme sensitivity compared with previous reports may have resulted from differences in contact area, direction of vibration, contact force, and the shape of the stimulus probe. The effects of each of these variables were studied. At 40 and 300 Hz (frequencies near the lower and upper end of the Pacinian range) thresholds were 9.8 and 18.5 dB (68 and 88%) lower, respectively, when subjects grasped the cylinder than when a 1-mm-diam probe vibrated perpendicular to the skin. These differences were accounted for as follows: 1) thresholds at a single fingerpad obtained with the large cylindrical surface were, on average, 20 and 60% lower, respectively, than thresholds with the punctate probe; 2) thresholds at the palm were, on average, 15 and 40% lower, respectively, than at the fingerpads; 3) thresholds obtained when the subjects grasped the cylinder averaged 40 and 20% less, respectively, than when the cylinder contacted only the palm; 4) thresholds with the cylinder contacting two fingers were 10 and 30% lower, respectively, than thresholds with the cylinder contacting a single finger; and 5) thresholds with vibration parallel to the skin surface were, on average, 10 and 30% lower, respectively, than thresholds with vibration perpendicular to the skin. Contact force, which was varied from 0.05 to 1.0 N, had no effect.

Hand↗
PubMed1999

SA1 and RA receptive fields, response variability, and population responses mapped with a probe array.

Twenty-four slowly adapting type 1 (SA1) and 26 rapidly adapting (RA) cutaneous mechanoreceptive afferents in the rhesus monkey were studied with an array of independently controlled, punctate probes that covered an entire fingerpad. Each afferent had a receptive field (RF) on a single fingerpad and was studied at 73 skin sites (50 mm2). The entire array was lowered to 1.6 mm below the point of initial skin contact (the background indentation) before delivering single-probe indentations. SA1 and RA responses differed in several ways. 1) SA1 RF boundaries were affected much less by indentation depth than were RA boundaries, and the SA1 RF areas were much more uniform in size. The mean SA1 RF area grew from 5.1 to 8.8 mm2 as the indentation depth increased from 50 to 500 microm; the mean RA RF area grew from 5.5 to 22.4 mm2 over the same intensity range. 2) SA1 RFs were more elongated than RA RFs. Elongated RFs were oriented in all directions relative to the skin ridges and the finger axis. 3) SA1 impulse rates were linear functions of indentation depth at all probe locations in the RF; RA responses tended toward saturation beginning at 100 microm indentation depth when the probe was over the HS. Similarities between SA1 and RA responses were that 1) both were extremely repeatable with SDs < 1 impulse per trial and 2) both had population responses (number of impulses) that were nearly linear functions of indentation depth. However, SA1s represented increasing indentation depth by increasing impulse rates in a small, relatively constant group of afferents, whereas the RAs represented increasing indentation depth predominantly by the recruitment of new afferents at a distance.

Animals↗
PubMed1999

Surround suppression in the responses of primate SA1 and RA mechanoreceptive afferents mapped with a probe array.

Twenty-four slowly adapting type 1 (SA1) and 26 rapidly adapting (RA) cutaneous mechanoreceptive afferents in the rhesus monkey were studied with an array of independently controlled, punctate probes that covered an entire fingerpad. Each afferent had a receptive field (RF) on a single fingerpad and was studied at 73 skin sites (50 mm2). The entire array was lowered to 1.6 to 3.0 mm below the point of initial skin contact (the background indentation) before delivering indentations with one to seven probes. Indentations were generally limited to 100 microm to minimize gross mechanical interactions. There were two major, new findings. 1) The discharge rates of both SA1 and RA afferents were strongly affected by the number of probes indenting the RF simultaneously. The effect was exponential. Each increase in probe number reduced the response by 24% in SA1 and 12% in RA afferents on average. When seven probes indented the skin simultaneously, the impulse rates in SA1 and RA afferents were reduced to 20 and 40% of the rates evoked by a single probe at the hot spot (all indentations were 100 microm). This shows that before any synaptic interaction in the CNS there is already a mechanism analogous to surround inhibition that suppresses an afferent's responses to uniform indentation and makes it especially sensitive to deviations from spatial uniformity. 2) The responses of both SA1 and RA afferents were independent of background array depth over the range from 1.6 to 3 mm below the point of initial skin contact. This shows that the neural responses to elements raised above a background are independent of the applied force over a wide range of forces. To relate the background depths to indentation force and to compare humans and monkeys, we studied the biomechanics of indentation with a uniform surface. A remarkable result is that the force-displacement relationships in humans and monkeys were the same; the skin is highly compliant for the first 2-3 mm of indentation and then becomes much stiffer. The results were the same in alert humans and monkeys and in monkeys anesthetized with pentobarbital. Ketamine anesthesia made the skin much stiffer and reduced the compliant range substantially.

Animals↗
PubMed1998

Structure of receptive fields in area 3b of primary somatosensory cortex in the alert monkey.

We investigated the two-dimensional structure of area 3b neuronal receptive fields (RFs) in three alert monkeys. Three hundred thirty neurons with RFs on the distal fingerpads were studied with scanned, random dot stimuli. Each neuron was stimulated continuously for 14 min, yielding 20,000 response data points. Excitatory and inhibitory components of each RF were determined with a modified linear regression algorithm. Analyses assessing goodness-of-fit, repeatability, and generality of the RFs were developed. Two hundred forty-seven neurons yielded highly repeatable RF estimates, and most RFs accounted for a large fraction of the explainable response of each neuron. Although the area 3b RF structures appeared to be continuously distributed, certain structural generalities were apparent. Most RFs (94%) contained a single, central region of excitation and one or more regions of inhibition located on one, two, three, or all four sides of the excitatory center. The shape, area, and strength of excitatory and inhibitory RF regions ranged widely. Half the RFs contained almost evenly balanced excitation and inhibition. The findings indicate that area 3b neurons act as local spatiotemporal filters that are maximally excited by the presence of particular stimulus features. We believe that form and texture perception are based on high-level representations and that area 3b is an intermediate stage in the processes leading to these representations. Two possibilities are considered: (1) that these high-level representations are basically somatotopic and that area 3b neurons amplify some features and suppress others, or (2) that these representations are highly transformed and that area 3b effects a step in the transformation.

Animals↗
PubMed1997

Neural coding mechanisms in tactile pattern recognition: the relative contributions of slowly and rapidly adapting mechanoreceptors to perceived roughness.

Tactile pattern recognition depends on form and texture perception. A principal dimension of texture perception is roughness, the neural coding of which was the focus of this study. Previous studies have shown that perceived roughness is not based on neural activity in the Pacinian or cutaneous slowly adapting type II (SAII) neural responses or on mean impulse rate or temporal patterning in the cutaneous slowly adapting type I (SAI) or rapidly adapting (RA) discharge evoked by a textured surface. However, those studies found very high correlations between roughness scaling by humans and measures of spatial variation in SAI and RA firing rates. The present study used textured surfaces composed of dots of varying height (280-620 micron) and diameter (0.25-2.5 mm) in psychophysical and neurophysiological experiments. RA responses were affected least by the range of dot diameters and heights that produced the widest variation in perceived roughness, and these responses could not account for the psychophysical data. In contrast, spatial variation in SAI impulse rate was correlated closely with perceived roughness over the whole stimulus range, and a single measure of SAI spatial variation accounts for the psychophysical data in this (0.974 correlation) and two previous studies. Analyses based on the possibility that perceived roughness depends on both afferent types suggest that if the RA response plays a role in roughness perception, it is one of mild inhibition. These data reinforce the hypothesis that SAI afferents are mainly responsible for information about form and texture whereas RA afferents are mainly responsible for information about flutter, slip, and motion across the skin surface.

Adaptation, Physiological↗
PubMed1997

Monkey cutaneous SAI and RA responses to raised and depressed scanned patterns: effects of width, height, orientation, and a raised surround.

Monkey cutaneous SAI and RA responses to raised and depressed scanned patterns: effects of width, height, orientation, and a raised surround. J. Neurophysiol. 78: 2503-2517, 1997. The aim of this study was to examine the slowly adapting type I (SAI) and rapidly adapting (RA) primary afferent representation of raised and depressed surface features. Isolated, raised, and depressed squares and small raised squares with a circular surround were scanned across the receptive fields of SAI and RA mechanoreceptive afferents innervating the distal fingerpads of the rhesus monkey. Pattern height ranged from -620 to +620 micron and width ranged from 0.2 to 7.0 mm. The surround radii ranged from 3.0 to 7.0 mm. Previous combined psychophysical and neurophysiological studies have provided evidence that SAI afferent responses are responsible for the perception of spatial form and texture and that RA afferents are responsible for the detection of stimuli that produce minute skin motion (flutter, slip, microgeometric surface features). Our results strengthen these hypotheses. Response properties shared by both SAI and RA afferent types were that both responded only to the edges of the larger raised and depressed patterns, both responded to falling edges half as vigorously as to rising edges, both responded to rising and falling edges with impulse rates that were proportional to the sine of the angle between the edge and the scanning direction, and both had suppressed responses to a small raised surface feature when a raised surround was closer than 6 mm. Response differences consistent with the hypothesis that SAI afferents are specialized for the representation of form were that SAI responses were confined to areas around the features that evoked them in areas that were 40-50% smaller than the comparable RA response areas, SAI responses were more than four times more sensitive to stimulus height than were RA afferents over the range from 280 to 620 micron, and SAI (but not RA) afferents responded 20-50% more vigorously to corners than to edges. Response differences consistent with the hypothesis that RA afferents are specialized for the detection of minute surfaces features were that only RA afferents responded to very small surface depressions, depressed squares 0.8 mm wide, that were detectable by palpation. Mechanisms underlying the many differences in SAI and RA response properties are discussed.

Action Potentials↗
PubMed1996

Marking microelectrode penetrations with fluorescent dyes.

Fluorescent dyes were used to mark and identify the tracks left by extracellular microelectrodes in neurophysiological experiments. Forty-two penetrations were made into the postcentral gyrus of 3 Macaque monkeys with electrodes coated with 1 of 5 fluorescent dyes (DiI, DiO, DiI-C5, PyPO, and Fast Blue). The electrodes were driven at rates ranging from 10 to 1000 microns/min, to a depth of about 4000 microns, where a small electrolytic lesion was made. Histological sections were viewed under fluorescent optics and the electrode tracks were reconstructed from the dye traces. Fluorescent traces (width 50-400 microns) were observed in 41 of 42 penetrations with 24 traces extending to the lesion site. Of the electrodes driven in less than 3 h, those coated with DiI (8/8) and DiI-C5 (8/8) left a trace to the lesion site, while 57% (4/7) of the DiO, 40% (2/5) of the Fast Blue and only 11% (1/9) of the PyPO tracks were fully marked. This method of marking penetrations can be used with any extracellular recording configuration, does not require tissue sections to be processed or stained, does not require electrical lesions, and causes no detectable tissue damage. Because the dyes fluoresce at different wavelengths, closely spaced tracks can be uniquely identified.

Animals↗
PubMed1995

Perspectives on the role of afferent signals in control of motor neuroprostheses.

K.O. Johnson reviews the architecture and low level neural mechanisms by which the external environment is transduced and encoded into the neural system, summarizing work that correlates neurophysiological and psychophysical testing with isolation of sensory components. The slowly adapting Type I afferent system is responsible for form and texture perception; the rapidly adapting afferent system is responsible for motion perception; and the Pacinian corpuscle system is responsible for vibratory sensation. R.R. Riso reviews the current level of understanding of the major factors to be considered in the design of a functional neuromuscular stimulation (FNS) grasp controller that uses cutaneous sensory feedback to detect slip. The elegant natural control scheme that matches the ratio of grip and lift forces to frictional conditions provides a model for implementing a slip-based control algorithm. D. Popović discusses the possible use of recordings from more proximal peripheral nerves to determine needed information for synthesis of locomotion. The discussion is illustrated with an animal model where rule-based closed-loop control is used for the ankle joint during treadmill locomotion. Neural signals from the tibial and superficial peroneal nerves were employed to substitute for missing afferent input from cutaneous and proprioceptive sensors. The feasibility of more invasive intraneural electrodes for distinguishing sensory from motor information in mixed nerves is considered. M. Koris raises surgical and functional issues relevant to developing clinical FNS systems. C. Van Doren suggests alternative neurophysiological and engineering approaches.

Afferent Pathways↗