Management of humeral nonunion after the failure of locking intramedullary nails.
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Biomedical subjects
Publications and source records attributed to K O Johnson.
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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.
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.
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.
Twenty-four subjects were studied before and up to 1 year after surgery that produced injury to a major sensory branch of the trigeminal nerve. We employed a battery of 11 psychophysical tests, in which the neural mechanisms underlying performance are understood, to study the basis of recovery following nerve injury. Immediately after nerve injury, sensation was profoundly impaired in all subjects. In the following weeks and months, the recovery of performance proceeded in an orderly fashion. Although the rates of recovery varied between subjects, the order of recovery between tasks did not. The recovery rates fell into three distinct categories. Recovery in one task, brush-stroke directional discrimination, was most rapid. Two weeks after nerve injury, 52% of subjects could discriminate brush-stroke direction; by 3 months only one subject could not perform this task. The second category comprised recovery rates for pain thresholds for noxious heat, cold and mechanical stimuli, and to preinjury performance in tasks assessing touch and vibration detection, two-point discrimination, cooling detection and subjective magnitude estimation of mechanical force. The third, slowest group included recovery rates for warming detection and grating orientation discrimination. Early recovery to preinjury performance levels in the brush-stroke direction and one-point versus two-point discrimination tasks was correlated with later recovery to near normal performance in the grating orientation task. The grating orientation task was unique in providing a measure that corresponded consistently with the subjects' reports of sensory deficits. Our psychophysical findings are consistent with neurophysiological data showing that the major primary afferent fibre classes reinnervate the skin at a similar rate. A hypothesis that accounts for the psychophysical findings in this study is that differences in recovery rates between tasks is determined largely by their relative dependencies on functional innervation density. Alternative hypotheses are considered.
Previous studies of the neural mechanisms of roughness perception have provided evidence that the magnitude of perceived roughness depends on spatial variation in the afferent population discharge. This paper reviews those studies and provides new data that appear to show that roughness perception cannot depend on activity in cutaneous rapidly adapting afferent fibers when surface element spacings exceed 1 mm. Finer surfaces have not been studied in combined psychophysical and neurophysiological studies. This paper also reinterprets the data of an earlier study, showing that all within-fiber neural coding mechanisms, which include impulse rate codes and codes based on the temporal structure of the afferent signal, can be rejected as the basis for roughness perception when the finger scans a textured surface.
The sensory neural pathways serving the lip, tongue, and finger are specialized for spatial information processing; thus, damage to these pathways is likely to be manifested most prominently as a loss of spatial acuity. For that reason, accurate measurement of spatial resolution at these regions is particularly important. The conventional test, the two-point discrimination task, does not measure the limit of spatial resolution and it yields variable results because it does not control nonspatial cues. The aim of this study was to quantify the limits of spatial resolution at the lip, tongue, and finger and to study the repeatability of those measurements using a stimulus that does not introduce nonspatial cues. We employed a grating orientation discrimination test, which has been studied extensively in relation to the underlying neural mechanisms. We obtained psychophysical thresholds for tactile spatial resolution from 15 normal, young adult subjects over seven test sessions. The finest gratings whose orientations were discriminated reliably had groove widths (gratings had equal groove and bar widths) that averaged 0.51 mm at the lip, 0.58 mm at the tongue, and 0.94 mm at the finger. These threshold measurements were highly reproducible between sessions with an overall improvement of 2% per session. These data suggest that the grating orientation discrimination task provides a stable, reliable measure of the human capacity for spatial resolution.
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Roughness perception is coded in the somatosensory system by neurons in the type I slowly adapting (SAI) system. When the fingers scan a surface, an isomorphic representation of the surface is encoded in the discharge patterns of SAI afferents. Central neurons in area 3b of primary somatosensory (SI) cortex spatially filter the peripheral image to compute local spatial variation. The outputs from these neurons converge onto neurons in area 1 and onto neurons in secondary somatosensory (SII) cortex which we believe is the critical processing pathway underlying roughness perception.
The effects of noise autocorrelation on neural waveform recognition (detection, classification, and superposition resolution) are investigated in this study using microelectrode recordings from the cortex of a monkey. Optimal waveform recognition is accomplished by passing the data through a whitening filter before matched filtering for detection or template matching for classification and superposition resolution. Template matching without whitening requires about 40% higher signal-to-noise ratio than template matching with whitening for comparable classification and superposition resolution. The comparable difference for detection is 15%.
1. The effects of selective attention were studied in SI and SII cortex of a rhesus monkey trained to perform two tasks, a tactile discrimination task and a visual detection task. In the tactile task, a letter was displayed on a video screen in front of the monkey and the animal was rewarded for responding when the raised letter (6.0 mm letter height) scanning across its finger (15 mm/s) matched the letter on the screen. In the visual task, three illuminated squares were displayed on the screen, and the animal was rewarded for detecting when one of the squares dimmed. The neural responses evoked by the raised letters were recorded continuously while the animal's focus of attention was switched back and forth between the two tasks. 2. Significant differences between the discharge rates evoked by raised letters in the two tasks were observed in approximately 50% of neurons in SI cortex and 80% of neurons in SII cortex. The effects in SII cortex were divided between increased (58%) and decreased (22%) rates. In SI cortex only increased rates were observed. 3. The attentional effects were expressed not only as changes in overall neuronal activity but also as modifications of the form of the responses evoked by the letters. 4. Whether attentional effects were observed depended upon the behavioral relevance of individual letters. During brief periods in the tactile task when a behavioral response could not yield a reward (time-out and reward periods) the neuronal responses were not significantly different from the responses evoked by the same letters during the visual task.(ABSTRACT TRUNCATED AT 250 WORDS)
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In the last decade or so, there has been rapid movement toward the use of more complex stimuli in the study of perceptual function related to the hand. This review has focused on the neural mechanisms of form and texture perception. Evidence from neurophysiological and psychophysical studies in which static touch, scanning touch, and the Optacon were used indicate that the spatial acuity of the RA system may be as much as three times poorer than the SAI system, evidence that suggests that form perception is dominated by the SAI system. Pattern recognition behavior in a tactual letter recognition task appears to be directly related to the response properties of SAI afferent fibers. Psychophysical studies of roughness perception show that roughness magnitude is related to surface structure in an orderly manner. Because roughness varies along an intensive continuum, mean impulse rate in one or more of the afferent systems is the most obvious coding possibility. No satisfactory relationship between mean impulse rate and roughness has been observed, however. The strongest hypothesis is that tactual roughness perception is based on spatial variation in the SAI population response. The combined evidence from studies reviewed here suggests complementary roles for each of the afferent systems, which are presented as working hypotheses: The SAI system is the primary spatial system and is responsible for tactual form and roughness perception when the fingers contact a surface directly and for the perception of external events through the distribution of forces across the skin surface. The PC system is responsible for the perception of external events that are manifested through transmitted high-frequency vibrations of the kind that are critical in the use of objects as tools. The RA system is responsible for the detection and representation of localized movement between skin and a surface as well as for surface form and texture when surface variation is too small to activate the SAI afferents effectively.
A previous study showed that roughness perception may depend on either temporal or spatial variations in firing rate among cutaneous mechanoreceptive afferents. The present study was designed to distinguish between these hypotheses. Plastic surfaces embossed with patterns of dots designed to produce predictable alterations in temporal and spatial firing rate variation were used as stimuli in psychophysical and neurophysiological experiments. Subjective roughness magnitudes obtained from psychophysical experiments fitted the predictions of the spatial but not the temporal hypothesis. In the neurophysiological experiments, the stimuli were scanned across the receptive fields of cutaneous mechanoreceptive afferents. Firing rate variation in the neural responses was measured using a range of temporal and spatial filters. Temporal variation was not correlated with roughness magnitude. Spatial variation, on a scale of 1-2 mm (one to two receptor spacings), was closely correlated with roughness.
The spatial resolving capacities of the four classes of mechanoreceptive afferents innervating human fingerpad skin were investigated to determine which class sets the limit of tactile spatial resolution for scanning stimuli. The stimulus consisted of an array of embossed dots (0.7 mm diameter, 0.5 mm high) arranged in a tetragonal pattern with dot spacing decreasing linearly from 6.4 mm at one end of the array to 0.87 mm at the other. The pattern was wrapped around a drum and repeatedly scanned across the receptive field of single afferents by continuously rotating the drum. Responses to many closely spaced scans were obtained by imposing a lateral shift of the pattern between each revolution. Impulses were recorded microneurographically. Responses were plotted in raster form to produce a neural image of the pattern. Responses of rapidly and slowly adapting type I (FAI and SAI) afferents resolved dots down to a spacing of about 1.5 mm. Responses of type II (FAII and SAII) afferents resolved dots down to a spacing of about 3.5 mm. Variation in scanning speed (range, 20-90 mm/sec) and contact force (range, 0.4-1.0 N) had minimal effects on spatial resolution of all afferents. The response clusters associated with individual widely spaced dots were used to investigate receptive field structure. FAI and SAI fields (mean areas, 6.1 and 4.8 mm2, respectively) each contained several zones of maximal sensitivity. FAI fields had five to eight such zones, whereas SAI fields had three to five such zones. As dot spacing decreased, neighboring dots interacted to affect the responses associated with the individual zones within a field. Initially, one or more zones were deactivated, effectively reducing receptive field size and allowing representation of finer spatial detail than would be predicted from the overall area of the receptive field. At very close dot spacings responses were only obtained when more than one sensitive zone within a field were simultaneously activated by different dots.
1. Subjects without any previous experience in a tactile psychophysics task participated in a study of tactile letter recognition employing active and passive touch. In the active task, subjects reached through a curtain and examined embossed letters with horizontal, unidirectional finger strokes. In the passive task, subjects sat with their arms and hands immobilized while a rotating drum stimulator pressed the embossed letters onto the right index finger. The stimulus conditions in the passive task were identical to those used in neurophysiological experiments with monkeys. 2. A survey of 40 naive subjects who were not screened in any way showed a wide range of performance levels. There was no difference between the subjects in the active and passive tasks, either in overall mean percent correct scores, which were 49.0 and 50.7%, respectively or in the percent correct scores for individual letters whose product-moment correlation coefficient was 0.94. The active and passive groups, which contained 25 and 15 members, respectively, had no members in common. 3. Videotapes of the finger movements of eight subjects in the active task showed a characteristic V-shaped velocity profile (velocity vs. lateral position) starting at approximately 100 mm/s at the left-hand edge of the plate containing the embossed letter, decelerating to a minimum when the center of the finger was directly over the letter, and then accelerating away from the letter. The average minimum scanning velocity was 17 mm/s. 4. Scanning velocity had no significant effect on performance in the passive task between 20 and 40 mm/s. An increase to 80 mm/s produced a 16% decline in percent correct identifications. 5. Learning effects were evident across sessions even though subjects were given no feedback or training. The increase in mean percent correct judgments averaged 4% per session, which lasted for approximately 1 h. 6. Data from 64 subjects were pooled for detailed comparison of identification patterns in active and passive touch. The results were analyzed and found to be consistent with the hypothesis that the identification and confusion probabilities are identical in the two modes. We conclude that there is no difference between active and passive touch in form recognition when the stimulus pattern is smaller than a finger pad. 7. Data from all experiments were pooled to produce a single confusion matrix with 324 presentations per letter. The majority of erroneous responses are grouped in a small number of confusion pairs and the majority of those confusion pairs are strongly asymmetric. The probable neural mechanisms of some confusion patterns are discussed.
We have used microneurographic techniques in human subjects to record mechanoreceptive afferent responses to standard Braille characters scanned across the finger pads. Responses from all four mechanoreceptor classes (FA I, FA II, SA I and SA II) have been reconstructed to form two-dimensional Spatial Event Plots (raster plots) of the Braille alphabet. Both the SA I and FA I responses resolve the dot patterns of Braille characters with sufficient fidelity that the representations of the individual characters can be recognized visually. Responses from SA II and FA II afferents do not resolve the Braille characters. We believe that the Spatial Event Plots derived from SA I and FA I responses closely approximate the images that are transmitted within a human nerve during Braille reading.
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