Stanley J. Bolanowski

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.

The Psychophysics of Tactile Perception and its Peripheral Physiological Basis

Publisher Summary This chapter examines the fundamental aspects of tactile psychophysics. The chapter also reviews the role of mechanoreceptive afferent neurons in mediating the psychophysical phenomena. This has proven to be a particularly fruitful avenue during the last two decades of research. The chapter focuses on tactile perceptions that are derived from the information provided by mechanoreceptors innervating the skin, and in some cases, subcutaneous tissues. Cutaneous mechanoreceptors are defined functionally as those elements of the peripheral nervous system (PNS) that are selectively responsive to non-noxious mechanical stimulation of the skin. A large body of data exists concerning the neurophysiological properties of cutaneous mechanoreceptors and their afferent fibers. This is reviewed in the chapter. The review encompasses the scientific advances over the last two decades related to tactile perception and its peripheral physiological basis. There are two significant features of this progress. First, the comparison between psychophysical and neurophysiological data is found to be valuable in experiments employing well-controlled and reproducible stimuli. Second, the development of more sophisticated models of somatosensation has provided greater insight into the neural basis of tactile perception, and has allowed for a wider range of experimental questions to be considered.

The frequency selectivity of information-processing channels in the tactile sensory system

The frequency selectivity of the P, NP I, and NP II channels of the four-channel model of mechanoreception for glabrous skin was measured psychophysically by an adaptation tuning curve procedure. The results substantially extend the frequency range over which the frequency selectivity of these channels is known and further confirm the hypothesis that the input stage of each of these channels consists of specific sensory nerve fibers and associated receptors. Specifically, the frequency characteristics of Pacinian nerve fibers, rapidly adapting (RA) nerve fibers, and slowly adapting Type II (SA II) nerve fibers were found to be the peripheral neurophysiological correlates of the P, NP I, and NP II channels, respectively. The finding that the tuning characteristic for a test stimulus of 250 Hz delivered through a small (0.008 cm2) contactor depended dramatically on the duration of the test stimulus whereas the detection threshold did not, provides new evidence in support of the hypothesis that separate NP II and P channels exist.

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.

Subjective magnitude of tactile roughness.

The subjective experience of tactile roughness was judged by subjects using the method of absolute magnitude estimation (AME). The stimuli were 11 grades of sandpaper having particle diameters ranging from 16 to 905 microm. All of the estimates resulted in power functions when assigned numbers were plotted as a function of particle diameter. It was determined that on the finger pad of the index finger and the thumb there was no difference between the active and passive modes of stimulation and that there was no difference in roughness estimates made on the finger and on the thumb. When the finger and thumb were stimulated simultaneously, higher numbers were assigned for a given stimulus indicating the presence of a form of spatial summation at these sites. The pleasantness of the tactile sensation, as assessed using AME, was inversely related to the roughness estimates. Furthermore, hydration of the stratum corneum with water and three concentrations of surfactant solutions reduced the sensation of roughness below that of normally hydrated skin.

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.

Effects of hydration on tactile sensation

The routine tasks of washing usually necessitates the immersion of parts of the body in water, which causes hydration and changes in the mechanical properties of the superficial layer of skin. To determine how hydration affects tactile sensations, the hydration and skin-surface temperature of glabrous and hairy skin was first measured under normal conditions (air), after submersion in distilled water alone and after submersion in a surfactant-water solution. In these experiments, measurements were made of the time to achieve complete hydration and the recovery time to normal levels. The uptake of water in hairy skin was found to be considerably greater than in glabrous skin, and retention was significantly prolonged by the surfactant additive. Subsequent experiments on glabrous skin, based on the results of the preceding hydration studies, measured in-air and hydrated tactile thresholds and sensation magnitudes to vibratory stimuli and to the roughness of textured surfaces. Vibrotactile detection thresholds were not affected by skin hydration, nor were sensation magnitudes to suprathreshold vibratory stimuli. However, suprathreshold perceptions of roughness were substantially altered by hydration. It is concluded that hydration and the mechanics of the skin play a major role in the perception of spatiotemporal (i.e., textured) surfaces and, thus, must be taken into account in any physiological/psychophysical model based on using such stimuli. This may not be required for models based on predominantly temporal (i.e., vibratory) stimuli.