Effects of receptor density on the tactile perception of roughness: implications for neural mechanisms of texture perception.

Aim of the study: The purpose of this study was to investigate the effects of receptor density in the glabrous skin of the hand on the perception of the roughness of a textured surface.Materials and methods: This was done by having observers make magnitude estimates of the perceived roughness of raised-dot surfaces at the fingertip, with its high receptor density, and the thenar eminence, with its much lower receptor density.Results: Judgments of perceived roughness averaged over the inter-dot spacings (0.8-5.9 mm) employed in the study did not differ significantly between the two sites, which suggested that roughness perception is not exclusively dependent upon a neural code involving variation in the activity levels of the nerve fibers of spatially distributed receptors, as is the case in spatial discrimination tasks such as spatial-gap detection, grove-orientation discrimination and letter recognition. This hypothesis was further supported by the finding that the elimination of temporal cues by preventing movement of the skin over the raised-dot surface drastically impaired judgments of perceived roughness at the thenar but had little effect on judgments of perceived roughness at the fingertip.Conclusion: These findings suggested that the neural code for perceived roughness at the fingertip is mediated primarily by spatial variation in the activity levels of spatially distributed receptors whereas the neural code for perceived roughness at the thenar is mediated primarily by temporal variation in the activity levels of individual receptors.

Perception of the tactile texture of raised-dot patterns: further evidence of an opponent process in the sense of touch.

Observers judged the tactile dissimilarities of raised-dot surfaces presented in pairs. The role of the SA I channel in determining these tactile dissimilarities was investigated by examining the dissimilarity judgments when this channel was adapted and when it was not. In an earlier study, the role of the PC channel in determining tactile dissimilarity was examined using the same stimulus materials when the PC channel was adapted and when it was not. Three orthogonal perceptual dimensions identified as blur, pattern roughness, and clarity were found using ALSCAL multidimensional analysis to account for the judged dissimilarities. The same three dimensions were found again in the present study. The dimensions of blur and pattern roughness were unaffected by adaptation of either the SA I or the PC channel. The finding of no effect of adaptation of the SA I channel on either of these two dimensions suggests that the roughness of the macrostructure of a textured surface is coded by the relative rather than by the absolute spatial variation in the firing rates of SA I nerve fibers. The dimension of dot clarity was strongly affected by adaptation of both the SA I channel and the PC channel. Adaptation of the PC channel increased dot clarity but adaptation of the SA I channel decreased it. This finding suggests that the perceived roughness of the microstructure of a textured surface is enhanced by the activity of the PC channel but decreased by the activity of the SA I channel.

Roughness perception in tactile channels: evidence for an opponent process in the sense of touch.

Magnitude estimates of the tactile roughness of raised-dot surfaces revealed that perceived overall roughness, defined as the combination of the perceived roughness of the dot pattern and the perceived roughness of the individual dots in the pattern, is an inverted U-shaped function of dot spacing, reaching a maximum at approximately 3.0 mm of dot separation. The hypothesis that Pacinian corpuscles are involved in roughness perception has been supported by the finding that selective adaptation of the Pacinian corpuscle (PC) channel with a 250-Hz stimulus at 20-dB SL results in a decrease in the perceived overall roughness of the raised-dot surface at the fingertip. The effect of PC channel adaptation on perceived overall roughness was attributable entirely to a reduction in the perceived roughness of the individual raised dots; PC adaptation had no effect on the perceived roughness of the raised-dot pattern. Selective adaptation of the slowly adapting type I (SA I) channel with a 5-Hz stimulus at 20-dB SL had the opposite effect of PC channel adaptation and resulted in an increase in the perceived roughness of the individual raised dots, and consequently the perceived overall roughness of the raised-dot surface. As was the case with PC channel adaptation, SA I channel adaptation had no effect on the perceived roughness of the pattern. Adaptation with a compound adapting stimulus containing 5- and 250-Hz components at 20-dB SL had no effect on perceived overall roughness, which suggests that the PC and SA I channels operate antagonistically in an opponent-process fashion in the perception of the microstructure of a textured surface. Neither PC adaptation nor SA I adaptation affected perceived pattern roughness, which suggests that pattern roughness is coded by relative rather than by absolute spatial variation in firing rate.

Learning in tactile channels.

Vibrotactile intensity-discrimination thresholds for sinusoidal stimuli applied to the thenar eminence of the hand declined as a function of practice. However, improvement was confined to the tactile information-processing channel in which learning had occurred. Specifically, improvements in performance with training within the Pacinian-corpuscle (PC) channel with a 250-Hz stimulus failed to transfer to performance within the rapidly adapting (RA) nerve fiber channel and the slowly adapting Type I (SA I) nerve fiber channel with a 20-Hz stimulus; similarly, improvements in performance with training within the RA/SA I channels failed to transfer to the PC channel. The hypothesis that tactile intensity-discrimination learning involves changes in sensory processes rather than the acquisition of the general skills required to perform the intensity-discrimination task is supported by the finding that improvements in performance with practice did not transfer to the untrained contralateral hand and by the finding of no transfer of training across channels. Within the PC channel, improvements in intensity discrimination with training transferred from the training intensity level (20-dB SL) to an untrained intensity level (10-dB SL), showing that learning within the PC channel under a specific stimulus condition can generalize to another stimulus condition provided both stimuli are processed by the PC channel. The finding that intensity-discrimination training, which resulted in a substantial reduction in the intensity difference limen, had no effect on the slope of the sensation-magnitude function suggests that tactile intensity-discrimination learning results primarily from a progressive reduction in neural noise over the course of training.

Perception of the tactile texture of raised-dot patterns: a multidimensional analysis.

An ALSCAL multidimensional scaling analysis in Euclidean space revealed that three orthogonal perceptual dimensions can account for the judged tactile dissimilarities of raised-dot patterns. Through magnitude estimates of various perceptual attributes, it was determined that the three dimensions consist of blur, roughness, and clarity. The only effect that selective adaptation of the Pacinian (P) channel had was to change the perceptual clarity of the raised dots against their background. Adaptation of the P channel with a 20 dB SL 250 Hz stimulus enhanced clarity. As indicated by magnitude estimates, adaptation of the P channel by the 250 Hz stimulus had no effect on the perceived roughness of the dot pattern but did cause the individual dots of the textured pattern to feel smoother. When the observer was required to estimate magnitude "overall roughness" defined as a combination of dot-pattern roughness and individual-dot roughness, adaptation of the P channel affected perceived roughness by reducing it. Taken as a whole, the results are consistent with the hypothesis that the NP channels and the P channel jointly influence the perception of textured surfaces.

Population-response model for vibrotactile spatial summation.

A computational model based on previous physiological and psychophysical data is presented for the human Pacinian (P) psychophysical channel. The model can predict the probability of detection in simple psychophysical tasks, and hence psychometric functions and thresholds. The model simulates stimulating variable and fixed glabrous skin sites with different-sized contactors and includes spatial variation of monkey P-fiber sensitivities. Therefore, it is especially suitable for studying spatial summation, i.e. the improvement of threshold with increasing contactor area. Selective contributions of neural integration (n.i.) and probability summation (p.s.) are also incorporated into the model. Model predictions are compared to psychophysical results of Gescheider et al. (2005). The performance of the model regarding the effects of contactor size is very good. In addition to predicting approximately 3 dB improvement of thresholds when the contactor area is doubled, the model also reveals nonlinear contributions of p.s. and n.i. Furthermore, the model asserts that thresholds are largely governed by neural integration when small contactors are used. These and other findings discussed in the article show that the presented model is a helpful tool for formulating testable hypotheses. Although the model can also simulate some temporal summation effects, simulation results do not conform well to previous data on temporal response properties. Thus, the model needs to be refined in that respect.

Spatial summation in the tactile sensory system: probability summation and neural integration.

Psychophysical thresholds for the detection of a 300-Hz burst of vibration applied to the thenar eminence were measured for stimuli applied to the skin through 1.5 cm2 and through 0.05 cm2 contactors. Thresholds were approximately 13 dB lower when the area of the contactor was 1.5 cm2 than when it was 0.05 cm2. The difference between the thresholds measured with the large and small contactors was significantly reduced when only the lowest thresholds obtained in the testing sessions were considered. This result supports the hypothesis that one component of spatial summation in the P channel is probability summation. In addition, threshold measurements within a session were less variable when measured with the 1.5 cm2 contactor. We conclude that spatial summation in the P channel is a joint function of two processes that occur as the areal extent of the stimulus increases: probability summation in which the probability of exceeding the psychophysical detection threshold increases as the number of receptors of varying sensitivities increases, and neural integration in which neural activity originating from separate receptors is combined within the central nervous system rendering the channel more sensitive to the stimulus.

Temporal gap detection in tactile channels.

The ability of observers to detect temporal gaps in bursts of sinusoids or bursts of band-limited noise was measured to assess the temporal acuity of Pacinian (P) and non-Pacinian (NP) tactile information processing channels. The P channel was isolated by delivering high frequency sinusoids or high frequency noise through a large 1.5-cm2 contactor to the thenar eminence. The NP channels were isolated from the P channel by delivering these stimuli as well as stimuli with lower frequencies through a small 0.01-cm2 contactor to the same site. Gap detection thresholds were higher for gaps in noise than for gaps in sinusoids but did not differ among conditions designed to isolate P and NP channels. The finding that temporal acuity does not differ among channels supports the hypothesis that, after termination of a stimulus, the P and NP channels exhibit the same amount of neural persistence. Also consistent with this hypothesis are the earlier findings that the enhancement of the sensation magnitude of a stimulus by a prior stimulus (Verrillo and Gescheider, Percept Psychophys 18: 128-136, 1975) and the duration of sensation after the termination of a stimulus (Gescheider et al., J Acoust Soc Am 91: 1690-1696, 1992) are independent of stimulus frequency. One important implication of this hypothesis, if true, is that the presence of temporal summation in the P channel and its absence in the NP channels, results, not from the lack of neural persistence in the NP channels, but instead, in marked contrast to the P channel, from the lack of a mechanism for integrating persistent neural activity over time.

Effect of aging on the subjective magnitude of vibration.

Two groups of subjects were tested using the method of Absolute Magnitude Estimation (AME) to determine the effect of age on the subjective intensity of vibration delivered to the skin of the hand. The mean age of the younger group was 23.5 years and that of the older group was 68.6 years. Average thresholds in the older group were higher in both the Pacinian (P) and non-Pacinian channel (NP II). The subjective magnitude of vibration was substantially lower at all intensities in the older group. Individual results clearly showed that the P channel saturates near the detection threshold of the NP II channel.

A four-channel analysis of the tactile sensitivity of the fingertip: frequency selectivity, spatial summation, and temporal summation.

Thresholds were measured for the detection of vibratory stimuli of variable frequency and duration applied to the index fingertip and thenar eminence through contactors of different sizes. The effects of stimulus frequency could be accounted for by the frequency characteristics of the Pacinian (P), non-Pacinian (NP) I, and NP III channels previously determined for the thenar eminence (Bolanowski et al., J Acoust Soc Am 84: 1680-1694, 1988; Gescheider et al., Somatosens Mot Res 18: 191-201, 2001). The effect of changing stimulus duration was also essentially identical for both sites, demonstrating the same amount of temporal summation in the P channel. Although the effect of changing stimulus frequency and changing stimulus duration did not differ for the two sites, the effect of varying the size of the stimulus was significantly greater for the thenar eminence than for the fingertip. The attenuated amount of spatial summation on the fingertip was interpreted as an indication that the mechanism of spatial summation consists of the operations of both neural integration and probability summation.