George A. Gescheider

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.

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.

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.

Some characteristics of tactile channels

The four information-processing channels of glabrous skin have distinct tuning characteristics which appear to be determined in the periphery at the level of sensory receptors and their afferent nerve fibers. The four-channel model [J Acoust Soc Am 84 (1988) 1680] has been updated to include measurement over a wider frequency range of tuning of the P and NP I channels, psychophysically determined by forward-masking and adaptation tuning curve methods. In addition to differences in their tuning, the P and NP channels differ in the following ways: (1) the P channel, but not NP channels, has been found to be capable of temporal summation, which operates by neural integration; (2) the capacity for spatial summation is also an exclusive property of the P channel; (3) sensitivity declines with age at a greater rate in the P channel than in the NP channels; (4) the masking or adaptation of a channel has no effect on the sensitivity of the other channels, although the channels interact in the summation of the perceived magnitudes of stimuli presented to separate channels.

Vibrotactile forward masking: Psychophysical evidence for a triplex theory of cutaneous mechanoreception

Threshold shifts for the detection of vibrotactile test stimuli were determined as a function of the intensity of a masker. A 50-ms sinusoidal test stimulus was applied to the thenar eminence of the hand 25 ms after the termination of a 700-ms sinusoidal masker applied to the same site. The frequency of the test stimulus and the frequency of the masker were varied. To eliminate the influence of the Pacinian receptor system, stimuli were delivered through a 0.01-cm2 contactor. The results support the hypothesis that the detection of vibration delivered through a small contactor is determined by two separate populations of non-Pacinian receptors. The study constitutes a psychophysical demonstration of the existence of three receptor systems responsible for the detection of vibration.

Stimulus context and absolute magnitude estimation: A study of individual differences

The effect of stimulus context on absolute-magnitude-estimation (AME) judgments was examined by determining whether the loudness judgment of a tone is influenced by the intensities of other tones presented within the session. A group of 18 subjects was tested in separate sessions in which they judged stimuli within either a low (10–60 dBSL) or a high (40–90 dBSL) range of intensities. Examination of the results of individual subjects revealed that judgments of stimuli common to the two ranges were, in most subjects, unaffected or only slightly affected by the position of the range. The judgments of 2 subjects who failed to follow the instructions, however, showed very large context effects due to changing the stimulus range. The results of a second experiment, in which 22 subjects judged the loudness of tones within either a narrow (35–65 dBSL) or a wide (20–80 dBSL)range, revealed that, in all but 1 subject, the width of the range had no systematic effect on the loudness judgments of stimuli common to both ranges. This was also true 1 month later when 16 of the subjects returned to the laboratory to judge the loudness of tones within an even wider range of 10–90 dBSL. It was concluded that AME judgments are relatively insensitive to the potential biasing influences of stimulus context.

Vibrotactile temporal summation for threshold and suprathreshold levels of stimulation

Threshold measurement and matching procedures were used to determine the amount of temporal summation at threshold and suprathreshold levels of vibrotactile stimulation on the thenar eminence of the hand. The frequency of the stimulus was 25, 40, 80, or 200 Hz. At 25 Hz, temporal summation was absent at all intensity levels. Considerable amounts of temporal summation were observed for 80- and 200-Hz stimuli, although the effects decreased as a function of intensity. At 40 Hz, no temporal summation was observed at threshold, but above threshold, a small amount was observed at all intensity levels. The results support a duplex model of mechanoreception in which one of two receptor systems exhibits temporal summation.

Enhancement and summation in the perception of two successive vibrotactile stimuli

Enhancement and summation were found to be fundamentally different perceptual processes affecting the sensation magnitude of two successive vibrotactile stimuli. Enhancement, defined operationally as an increment in the subjective magnitude of one stimulus due to the presentation of a prior stimulus, and summation, defined as an increment in overall subjective magnitude of the two stimuli, were measured for sinusoidal vibration of the thenar eminence of the hand. The effect of summation was maximum when the two stimuli greatly differed in frequency, whereas maximum enhancement effects were found when both stimuli were close in frequency. The summation effect showed little decay as the interstimulus interval was increased to as much as 500 msec, whereas enhancement effects decayed to zero at approximately 500 msec. Results were similar to those obtained in comparable studies of audition and support the hypothesis that there are at least two distinct information-processing channels for the perception of cutaneous vibration.

The measurement of loudness in individual children and adults by absolute magnitude estimation and cross‐modality matching

Twelve adults and 11 children (age range 4-7 years) performed absolute magnitude estimation of the apparent lengths of lines and the loudnesses of 1000-Hz tones as well as cross-modality matching between loudness and apparent line length. Consistent with the notion that children and adults have similar impressions of loudness, there were no major differences between the absolute magnitude estimation (AME) and cross-modality matching (CMM) data of the adults and children. A direct comparison between the exponents for loudness by AME and CMM was made when a correction factor was employed to eliminate the effects of idiosyncratic use of numbers from the AME exponents. The results support the hypothesis that, with proper instructions, both children and adults can judge stimuli on an absolute scale. Specifically, for 9 out of 12 adults and 9 out of 11 children, lines and tones assigned the same number in absolute magnitude estimation were judged to be subjectively equal in cross-modality matching.