Ronald T. Verrillo

Four channels mediate the mechanical aspects of touch

Although previous physiological and anatomical experiments have identified four afferent fiber types (PC, RA, SA II, and SA I) in glabrous (nonhairy) skin of the human somatosensory periphery, only three have been shown to mediate tactile (mechanoreceptive) sensation. Psychophysical evidence that four channels (P, NP I, NP II, and NP III) do, indeed, participate in the perceptual process is presented. In a series of experiments involving selective masking of the various channels, modification of the skin-surface temperature, and testing cutaneous sensitivity down to very low-vibratory frequencies, the fourth psychophysical channel (NP III) is defined. Based on these experiments and previous work from our laboratory, it is concluded that the four channels work in conjunction at threshold to create an operating range for the perception of vibration that extends from at least 0.4 to greater than 500 Hz. Each of the four channels appears to mediate specific portions of the overall threshold-frequency characteristic. Selection of appropriate neural-response criteria from previously published physiological data and correlation of their derived frequency characteristics with the four psychophysical channels indicates that each channel has its own physiological substrate: P channel and PC fibers, NP I channel and RA fibers, NP II channel and SA II fibers, and NP III channel and SA I fibers. These channels partially overlap in their absolute sensitivities, making it likely that suprathreshold stimuli may activate two or more of the channels at the same time. Thus the perceptual qualities of touch may be determined by the combined inputs from four channels.

Effect of Contactor Area on the Vibrotactile Threshold

Vibrotactile thresholds were determined as a function of frequency, contactor configuration, and contactor area. It was found that contactor area is a more important stimulus parameter than the gradient or curvature of displacement. The absolute threshold for vibration seems to be independent of frequency when very small contactors are used and independent of area at low frequencies. For higher values of these parameters, it strongly depends on both. It seems reasonable to conclude that responses to mechanical displacement of the skin are mediated by more than one receptor system, although direct evidence is still lacking.

Sensation magnitude of vibrotactile stimuli

A systematic investigation of the subjective magnitude of vibrotaction was undertaken to: (1) determine the growth of sensation as a function of stimulus intensity; (2) establish contours of equal subjective magnitude; and (3) compare over a wide range of frequency and intensity the psychophysical methods of direct scaling and intensity matching. The results show that the data obtained by direct scaling are comparable to the data obtained by interfrequency matching. The subjective magnitude function is a power function with a slope of about 0.89 for frequencies up to 350 Hz. Near threshold the growth of sensation is proportional to the physical intensity. Contours of equal subjective magnitude for vibration across 10 frequencies and at 11 levels of intensity are given.

Investigation of Some Parameters of the Cutaneous Threshold for Vibration

Sensitivity to vibration on the hand was determined as a function of frequency, contactor dimensions, contactor configuration, and distance of the contactor from a rigid support. It was found that each of these parameters affects the threshold in a different way. In the frequency range between 25 and 640 cps, the absolute threshold as a function of frequency yields a U‐shaped curve that reaches a maximum of sensitivity in the region of 250 cps. The effect of the geometric parameters appears to be highly complex.

The Effects of Aging on Information-Processing Channels in the Sense of Touch: I. Absolute Sensitivity

Thresholds for detecting vibrotactile signals of variable frequency applied to the thenar eminence of the hand by small and large contactors were measured in subjects ranging in age from 10 to 89 years. Thresholds were found to increase as a function of age, but the rate of increase was greater after than before the age of 65 years. The rate of loss of vibrotactile sensitivity was substantially greater in the P channel (mediated by Pacinian corpuscles) than in the NP I channel (mediated by rapidly adapting fibers), the NP II channel (mediated by slowly adapting type II fibers), or the NP III channel (mediated by slowly adapting type I fibers). Women were frequently found to have greater sensitivity than men.

Psychophysics of vibrotactile stimulation

The effects of a wide range of stimulus parameters upon the psychophysical responses of subjects are important to an understanding of the functional characteristics of neural systems that underlie responses to cutaneous mechanical stimulation and the interactions that may occur between these systems. This understanding may also be useful to design considerations in the development of devices for tactile communication. This paper begins with a brief description of the two major systems in terms of psychophysical measurements and then focuses on a more detailed examination of possible interactions between them. The effects of time and space variables, such as frequency, temporal and intensity relationships between stimuli, area of stimulation, and surface gradients on the skin, are considered.

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.

Hairy skin: psychophysical channels and their physiological substrates.

Experiments were conducted in which threshold-frequency characteristics were measured on the hairy skin of the forearm of human observers. Thresholds were measured with two stimulus probe areas (2.9 and 0.008 cm2) at three skin-surface temperatures (15 degrees, 30 degrees, and 40 degrees C). The results suggest that whereas glabrous skin uses four distinct channels of information, only three channels may be involved in mediating the sense of touch for hairy skin. The three channels are defined as Ph, (Pacinian, hairy skin), NPh low (non-Pacinian, hairy skin, low frequencies) and NPh mid (non-Pacinian, hairy skin, middle frequencies). In addition, it is proposed that the neural substrates for the three psychophysically characterized channels are, respectively, the Pacinian corpuscle (PC) nerve fibers, the slowly adapting type II (SAII) fibers, and the rapidly adapting (RA) fibers.

Vibrotacile masking of Pacinian and non‐Pacinian channels

Vibrotactile masking functions were determined using sinusoidal and noise maskers. Results were nearly identical within the Pacinian (P) and non-Pacinian (NP) channels. At low maskers SLs there was a substantial amount of negative masking which proved not be an artifact of stimulus definition. The critical parameters for successful prediction of the data were a peripheral threshold and internal Gaussian noise. Threshold shifts in cross-channel stimulation can be attributed to the masker exceeding the detection threshold of the signal channel.

The effects of skin temperature on the psychophysical responses to vibration on glabrous and hairy skin.

Psychophysical detection thresholds for vibration were measured at the thenar eminence and volar forearm using a 0.008-cm2 (1.0-mm-diam) contactor. Measurements were made at 14 sinusoidal frequencies between 12 and 500 Hz at six skin temperatures between 15 degrees and 40 degrees C. The results are consistent with the hypothesis that three functionally discrete non-Pacinian afferent systems mediate vibrotactile responses. It was possible to identify the response characteristic of the rapidly adapting (Meissner) system, but it was not possible to isolate the responses of two slowly adapting (SAI, SAII) systems.