Conditioned pain modulation dampens the thermal grill illusion.

BACKGROUND: The thermal grill illusion (TGI) refers to the perception of burning heat and often pain that arises from simultaneous cutaneous application of innocuous warm and cool stimuli. This study utilized conditioned pain modulation (CPM) to help elucidate the TGI's underlying neural mechanisms, including the debated role of ascending nociceptive signals in generating the illusion. METHODS: To trigger CPM, subjects placed the left hand in noxious cold (6 °C) water before placing the right volar forearm onto a thermal grill. Lower pain and unpleasantness ratings of the grill in this CPM run compared to those in a control run (i.e. 33 °C water) were taken as evidence of CPM. To determine whether CPM reduces noxious heat pain and illusory heat pain equally, an experimental group of subjects rated pain and unpleasantness of a grill consisting of innocuous alternating warm (42 °C) and cool (18 °C) bars, while a control group rated a grill with all bars controlled to a noxious temperature (45 °C). RESULTS: CPM produced significant and comparable reductions in pain, unpleasantness and perceived heat of both noxious heat and the TGI. CONCLUSIONS: This result suggests that the TGI results from signals in nociceptive dorsal horn convergent neurons, since CPM involves descending inhibition with high selectivity for this neuronal population. More broadly, CPM's ability to produce a shift in perceived thermal sensation of both noxious heat and the TGI from 'hot' to 'warm' implies that nociceptive signals generated by a cutaneous stimulus can contribute to its perceived thermal intensity. SIGNIFICANCE: Conditioned pain modulation reduces the perceived painfulness, unpleasantness and heat of the thermal grill illusion and noxious heat similarly. The results have important theoretical implications for both types of pain.

Field reversed configuration confinement enhancement through edge biasing and neutral beam injection.

Field reversed configurations (FRCs) with high confinement are obtained in the C-2 device by combining plasma gun edge biasing and neutral beam injection. The plasma gun creates an inward radial electric field that counters the usual FRC spin-up. The n = 2 rotational instability is stabilized without applying quadrupole magnetic fields. The FRCs are nearly axisymmetric, which enables fast ion confinement. The plasma gun also produces E × B shear in the FRC edge layer, which may explain the observed improved particle transport. The FRC confinement times are improved by factors 2 to 4, and the plasma lifetimes are extended from 1 to up to 4 ms.

Dynamic formation of a hot field reversed configuration with improved confinement by supersonic merging of two colliding high-ß compact toroids.

A hot stable field-reversed configuration (FRC) has been produced in the C-2 experiment by colliding and merging two high-ß plasmoids preformed by the dynamic version of field-reversed θ-pinch technology. The merging process exhibits the highest poloidal flux amplification obtained in a magnetic confinement system (over tenfold increase). Most of the kinetic energy is converted into thermal energy with total temperature (T{i}+T{e}) exceeding 0.5 keV. The final FRC state exhibits a record FRC lifetime with flux confinement approaching classical values. These findings should have significant implications for fusion research and the physics of magnetic reconnection.

Human vibrotactile frequency discriminative capacity after adaptation to 25 Hz or 200 Hz stimulation.

A two-interval forced-choice (2-IFC) tracking procedure was used to evaluate the effects of a 15-s pre-exposure to either 25 Hz or 200 Hz stimulation ("25 Hz or 200 Hz adaptation") on human vibrotactile frequency discrimination threshold (frequency DL/Weber fraction). Three subjects were studied. All stimuli (standard and comparison) were delivered to a central location on the thenar eminence of the hand. The frequency DL/Weber fraction was determined for each subject under the following conditions: (1) no recent prior exposure to vibrotactile stimulation ("unadapted"); (2) after 15 s adaptation to 25 Hz stimulation; and (3) after 15 s adaptation to 200 Hz stimulation. The results demonstrate that the effects of frequency of adaptation on frequency discriminative capacity when the standard stimulus is 25 Hz are not the same as when the standard stimulus is 200 Hz. The differential changes in the capacity of subjects to discriminate frequency of cutaneous flutter (10-50 Hz) or vibratory (>200 Hz) stimulation that occur subsequent to a 15-s exposure of the thenar to 25 Hz or 200 Hz stimulation are proposed to reflect frequency-specific, adaptation-induced modification of the response of contralateral primary somatosensory cortex (SI and SII) to skin mechanoreceptor afferent drive.

Vibrotaction and texture perception.

Several recent studies support Katz's hypothesis that vibrotaction plays a role in the perception of tactile textures with elements too small and closely spaced to be processed spatially. For example, eliminating vibration by preventing movement of a stimulus surface across the skin compromises psychophysical scaling and discrimination of fine, but not coarse, textures. Fine-texture discrimination is also impaired when vibrotactile channels are desensitized by adaptation. A role for vibrotaction in texture perception is plausible, given the keenness of this submodality: the sensory qualities produced by a sinusoidal vibration uniquely specify its frequency and amplitude, and subjects can distinguish some complex vibrations that differ in waveform but have the same spectral components. Finally, imposed vibration can modify the perceived texture of a haptically-examined surface. Taken together, these lines of evidence support the view that vibrotaction is both necessary and sufficient for the perception of fine tactile textures.

Local vibrotactile and pain sensitivities are negatively related in temporomandibular disorders.

Earlier research has shown that cutaneous experimental pain can elevate the vibrotactile threshold at the same skin locus. The purpose of this study was to determine whether vibrotactile and pain thresholds in a clinical (temporomandibular disorders [TMD]) population are consistent with the hypothesis that chronic pain causes a similar elevation. Specifically, we predicted that TMD subjects with soreness (low palpation-pain threshold) at a given skin site would have relatively high vibrotactile thresholds at the same location. Measurements on the skin overlying the masseter in 18 individuals with TMD showed that pain sensitivity was negatively correlated with sensitivity to 20-Hz vibration (presumed to activate a rapidly adapting mechanoreceptive channel), but not with sensitivity to 200-Hz vibration (thought to activate primarily a slowly adapting channel, because the Pacinian channel is lacking in the orofacial region). There was no relationship between vibration thresholds over the masseter and pain threshold at other orofacial sites, including the contralateral masseter. Vibrotactile and pain thresholds were uncorrelated in control participants without chronic pain (n = 18). The results indicate that in TMD, a localized relationship exists between pain sensitivity and the sensitivity of a low-frequency vibrotactile channel.

Vibrotactile adaptation impairs discrimination of fine, but not coarse, textures.

The effect of vibrotactile adaptation on the ability to discriminate textured surfaces was examined in three experiments. The surfaces were rectilinear arrays of pyramids produced by etching of silicon wafers. Adaptation to 100-Hz vibration severely hampered discrimination of surfaces with spatial periods below 100 microm (Experiment 1), but had little effect on the discrimination of coarser textures (Experiment 2). To determine which vibrotactile channel--Rapidly Adapting or Pacinian--plays the larger role in mediating the discrimination of fine textures, widely separated adapting frequencies (10 and 250 Hz) were used in Experiment 3. The fact that high- but not low-frequency adaptation interfered with discrimination suggests that the Pacinian system contributes importantly to this ability. Taken as a whole, the results of this study strongly support the duplex theory of tactile texture perception, according to which different mechanisms--spatial and vibrotactile--mediate the perception of coarse and fine textures, respectively.

Complex tactile waveform discrimination.

Complex vibrotactile waveforms consisting of two superimposed sinusoids at varying phases were presented to the fingertip, and observers made "same-different" judgments. It was found that the low-frequency (10Hz+30Hz) waveforms were discriminable from one another while discrimination of the high-frequency (100Hz+300Hz) vibrations was poor. High-frequency adaptation did not impair discrimination of the low-frequency waveforms, suggesting that the RA channel mediated discrimination. Low-frequency adaptation impaired discrimination of the high-frequency stimuli, suggesting that the RA channel likewise mediated the modest level of performance observed in the absence of an adapting stimulus. The results indicate that this channel encodes complex waveforms temporally. A simple model for low-frequency waveform discrimination is proposed. The results obtained with the high-frequency complex waveforms are compatible with the hypothesis that the PC channel integrates stimulus energy over time.

Evidence for the duplex theory of tactile texture perception.

Three experiments are reported bearing on Katz's hypothesis that tactile texture perception is mediated by vibrational cues in the case of fine textures and by spatial cues in the case of coarse textures. Psychophysical responses when abrasive surfaces moved across the skin were compared with those obtained during static touch, which does not provide vibrational cues. Experiment 1 used two-interval forced-choice procedures to measure discrimination of surfaces. Fine surfaces that were readily discriminated when moved across the skin became indistinguishable in the absence of movement; coarse surfaces, however, were equally discriminable in moving and stationary conditions. This was shown not to result from any inherently greater difficulty of fine-texture discrimination. Experiments 2 and 3 used free magnitude estimation to obtain a more comprehensive picture of the effect of movement on texture (roughness) perception. Without movement, perception was seriously degraded (the psychophysical magnitude function was flattened) for textures with element sizes below 100 microns; above this point, however, the elimination of movement produced an overall decrease in roughness, but not in the slope of the magnitude function. Thus, two components of stimulation (presumably vibrational and spatial) contribute to texture perception, as Katz maintained; mechanisms for responding to the latter appear to be engaged at texture element sizes down to 100 microns, a surprisingly small value.

Individual differences in perceptual space for tactile textures: evidence from multidimensional scaling.

Ratio scaling was used to obtain from 5 subjects estimates of the subjective dissimilarity between the members of all possible pairs of 17 tactile surfaces. The stimuli were a diverse array of everyday surfaces, such as corduroy, sandpaper, and synthetic fur. The results were analyzed using the multidimensional scaling (MDS) program ALSCAL. There was substantial, but not complete, agreement across subjects in the spatial arrangement of perceived textures. Scree plots and multivariate analysis suggested that, for some subjects, a two-dimensional space was the optimal MDS solution, whereas for other subjects, a three-dimensional space was indicated. Subsequent to their dissimilarity scaling, subjects rated each stimulus on each of five adjective scales. Consistent with earlier research, two of these (rough/smooth and soft/hard) were robustly related to the space for all subjects. A third scale, sticky/slippery, was more variably related to the dissimilarity data: regressed into three-dimensional MDS space, it was angled steeply into the third dimension only for subjects whose scree plots favored a nonplanar solution. We conclude that the sticky/slippery dimension is perceptually weighted less than the rough/smooth and soft/hard dimensions, materially contributing to the structure of perceptual space only in some individuals.

Imposed vibration influences perceived tactile smoothness.

According to the duplex theory of tactile texture perception, detection of cutaneous vibrations produced when the exploring finger moves across a surface contributes importantly to the perception of fine textures. If this is true, a vibrating surface should feel different from a stationary one. To test this prediction, experiments were conducted in which subjects examined two identical surfaces, one of which was surreptitiously made to vibrate, and judged which of the two was smoother. In experiment 1, the vibrating surface was less and less often judged smoother as the amplitude of (150 Hz) vibration increased. The effect was comparable in subjects who realized the surface was vibrating and those who did not. Experiment 2 showed that different frequencies (150-400 Hz) were equally effective in eliciting the effect when equated in sensation level (dB SL). The results suggest that vibrotaction contributes to texture perception, and that, at least within the Pacinian channel, it does so by means of an intensity code.

A ratio code for vibrotactile pitch.

Subjective impressions of pitch for 80 different sinusoidal vibrotactile stimuli delivered to the index finger were measured by free magnitude estimation in four subjects. In three of the subjects, pitch at a given frequency decreased as stimulus amplitude increased. The data of these subjects were well described by a model of pitch based on the relative levels of activation of the three major tactile channels. The main element in this model was a ratio of P channel activity to the sum of the activity levels of the P, NPI, and NPIII channels. Activity levels of the channels were estimated on the basis of the psychophysical literature, including a study of vibrotactile loudness using the same subjects and stimuli as those employed here. A fourth subject, whose pattern of loudness judgments had previously been shown to differ from those of the other subjects, did not conform to this pitch model: her data revealed significant increases in pitch with increases in amplitude, and appear to reflect an inability to combine signals across vibrotactile channels. Pitch changes resulting from vibrotactile adaptation were directionally consistent with our ratio model: pitch was slightly increased by adaptation to a 25 Hz stimulus, and slightly decreased by 200 Hz adaptation.

Adaptation-induced enhancement of vibrotactile amplitude discrimination: the role of adapting frequency.

Two-interval forced-choice tracking was used to measure amplitude discrimination for 20-Hz vibrotactile test stimuli presented to the thenar eminence of three human observers. For all observers, relative difference threshold could be decreased by adaptation to a 20- or 100-Hz stimulus. Maximal enhancement of discrimination occurred when the amplitude of the adapting stimulus was such that it excited the NP I system to approximately the same degree that the test stimuli did. A signal detection analysis determined that shifts in the observers' criteria could not consistently account for the enhancement of amplitude discrimination. A more likely explanation, in view of recent physiological discoveries, is that under optimal conditions of adaptation test stimuli differing slightly in amplitude become more distinctive because CNS events underlying the resultant sensory experiences become more refined and stimulus specific.

Perceived intensity of vibrotactile stimuli: the role of mechanoreceptive channels.

The perceived intensity of vibrotactile stimuli was studied by means of free magnitude estimation. Eighty different sinusoidal stimuli ranging in frequency from 10 to 200 Hz, and in amplitude from 2.4 to 154 microns, were presented to the left index fingerpad of psychophysical observers through a 5-mm-diameter contactor. Estimates at a given frequency increased with amplitude in all four subjects, and estimates at a given amplitude increased with frequency in three. For the fourth subject, however, intermediate frequencies (25-75 Hz) produced the most intense sensations; the relative sensory effectiveness of different frequencies suggested that in her case, perceived vibrotactile intensity was determined largely by signals in Meissner afferents. From the data of this unusual subject, and from high-frequency (200-Hz) measurements on the normal subjects, quantitative descriptions were derived of the signals in Meissner and Pacinian channels, respectively, that could contribute to subjective intensity. Candidate algorithms by which the signals from the two channels might interact were then evaluated by comparison of modeled and empirically determined subjective intensity values. It was found that subjective intensity is given by the sum of (1) the stronger of the two channels' signals, and (2) half the weaker signal, the latter apparently being reduced by cross-channel suppression that occurs only at suprathreshold levels. Adapting to 25-Hz vibration selectively reduces the perceived intensity of low frequencies, whereas adapting to 200-Hz vibration has a corresponding effect at high frequencies. It is concluded that an understanding of perceived vibrotactile intensity requires knowledge of the signals in vibrotactile channels, and of the interactions between those channels.

Vibrotactile adaptation enhances frequency discrimination.

Human vibrotactile frequency discrimination (with respect to a 25-Hz standard stimulus, 20 dB above unadapted detection threshold) was measured on the thenar eminence and index fingerpad, using two-interval forced-choice tracking. Measurements were made in the unadapted state and following exposure to 25-Hz adapting stimuli of various amplitudes. The standard and all comparison stimuli were equated for perceived intensity, on the basis of matching experiments that were carried out separately under each adapting condition. Frequency difference thresholds were lowest when the amplitude of the adapting stimulus was equal to the amplitude of the standard. This result complements the earlier finding [A. K. Goble and M. Hollins, J. Acoust. Soc. Am. 93, 418-424 (1993)] that adaptation sharpens amplitude discrimination of supraliminal stimuli that are similar to the adapting stimulus. Taken together, these discoveries suggest that somatosensory mechanisms that are engaged by extended stimulation serve to enhance detection of changes in the properties, both quantitative and qualitative, of that stimulation.

The tactile movement aftereffect.

The existence of a tactile movement aftereffect was established in a series of experiments on the palmar surface of the hand and fingers of psychophysical observers. During adaptation, observers cupped their hand around a moving drum for up to 3 min; following this period of stimulation, they typically reported an aftereffect consisting of movement sensations located on and deep to the skin, and lasting for up to 1 min. Preliminary experiments comparing a number of stimulus materials mounted on the drum demonstrated that a surface approximating a low-spatial-frequency square wave, with a smooth microtexture, was especially effective at inducing the aftereffect; this adapting stimulus was therefore used throughout the two main experiments. In Experiment 1, the vividness of the aftereffect produced by 2 min of adaptation was determined under three test conditions: with the hand (1) remaining on the now stationary drum; (2) in contact with a soft, textured surface; or (3) suspended in air. Subjects' free magnitude estimates of the peak vividness of the aftereffect were not significantly different across conditions; each subject experienced the aftereffect at least once under each condition. Thus the tactile movement aftereffect does not seem to depend critically on the ponditions of stimulation that obtain while it is being experienced. In Experiment 2, the vividness and duration of the aftereffect were measured as a function of the duration of the adapting stimulus. Both measures increased steadily over the range of durations explored (30-180 sec). In its dependence on adapting duration, the aftereffect resembles the waterfall illusion in vision. An explanation for the tactile movement aftereffect is proposed, based on the model of cortical dynamics of Whitsel et al. (1989, 1991). With assumed modest variation of one parameter across individuals, this application of the model is able to account both for the data of the majority of subjects, who experienced the aftereffect as opposite in direction to the adapting stimulus, and for those of an anomalous subject, who consistently experienced the aftereffect as being in the same direction as the adapting stimulus.

Perceptual dimensions of tactile surface texture: a multidimensional scaling analysis.

The purpose of this study was to examine the subjective dimensionality of tactile surface texture perception. Seventeen tactile stimuli, such as wood, sandpaper, and velvet, were moved across the index finger of the subject, who sorted them into categories on the basis of perceived similarity. Multidimensional scaling (MDS) techniques were then used to position the stimuli in a perceptual space on the basis of combined data of 20 subjects. A three-dimensional space was judged to give a satisfactory representation of the data. Subjects' ratings of each stimulus on five scales representing putative dimensions of perceived surface texture were then fitted by regression analysis into the MDS space. Roughness-smoothness and hardness-softness were found to be robust and orthogonal dimensions; the third dimension did not correspond closely with any of the rating scales used, but post hoc inspection of the data suggested that it may reflect the compressional elasticity ("springiness") of the surface.

Vibrotactile adaptation enhances amplitude discrimination.

Human psychophysical detection and amplitude discrimination thresholds for 25-Hz sinusoidal vibrations were measured on the thenar eminence using two-interval forced-choice tracking, in the unadapted state and following exposure to 25-Hz adapting stimuli representing a range of amplitudes (5-25 dB SL). As expected, detection threshold was elevated 6 to 7 dB for each 10-dB increase in the adapting stimulus. In contrast, amplitude difference thresholds for 10 and 20 dB SL standard stimuli were generally lowest when the amplitude of the adapting stimulus was equal to the amplitude of the standard. The results indicate that while adaptation impairs detection of a liminal vibrotactile stimulus, it improves intensity discrimination of supraliminal stimuli that are close in amplitude to the adapting stimulus. The compatability between these results and a recently proposed model of cortical dynamics (Whitsel et al., 1989) suggests that cortical events may contribute significantly to the physiological basis of vibrotactile adaptation.

Vibrotactile adaptation on the face.

Threshold amplitude for vibration is elevated if testing is preceded by extended exposure to a vibratory adapting stimulus of appropriate amplitude and frequency. This phenomenon, previously studied almost exclusively on the hand, is here shown for the first time to occur on the face as well. Adaptation is then used analytically to determine that the two-branched threshold-versus-frequency function obtained on the face by Verrillo and Ecker (1977) represents the activity of two distinct mechanisms. Action spectra of vibrotactile adaptation reveal the presence of both mechanisms even in subjects whose unadapted threshold function (like that reported by Barlow, 1987) shows no sign of duplexity. Finally, the data suggest that on the face (unlike the hand), cross-channel adaptation may occur at high adapting amplitudes.