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.     

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.     

Thresholds for the perception of hand-transmitted vibration: dependence on contact area and contact location

The detection of vibration applied to the glabrous skin of the hand varies with contact conditions. Three experiments have been conducted to relate variations in the perception of hand-transmitted vibration to previously reported properties of tactile channels. The effects of a surround around the area of contact, the size of the area of contact, the location of the area of contact, the contact force, and the hand posture on perception of thresholds were determined for 8-500 Hz vibration. Removal of a surround around a contact area on the fingertip elevated thresholds of the NP II channel (FA I fibres) at frequencies less than 31.5 Hz and reduced thresholds of the Pacinian channel (FA II fibres) at frequencies greater than about 63 Hz. When no surround was present, thresholds reduced systematically as the contact area increased from the fingertip to the whole hand at frequencies from 16 to 125 Hz, although the decrease was not inversely proportional to the increase in contact area. The results are partly explained by spatial summation in the Pacinian channel (FA II fibres) and the involvement of the NP II channel (SA II) with some influence of biodynamic responses and contact pressures. There were regional differences in sensitivity over the hand within the NP I channel but not within the Pacinian channel: the NP I thresholds (less than 31.5 Hz) decreased from proximal to distal regions of the hand, whereas the Pacinian thresholds (125 Hz) were independent of contact location over the hand.     

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.     

Independent responses of Pacinian and Non-Pacinian systems with hand-transmitted vibration detected from masked thresholds

This study was designed to identify psychophysical channels responsible for the detection of hand-transmitted vibration. Perception thresholds for vibration (16, 31.5, 63 and 125?Hz sinusoidal for 600?ms) at the distal phalanx of the middle finger and the whole hand were determined with and without simultaneous masking stimuli (1/3 octave bandwidth Gaussian random vibration centered on either 16?Hz or 125?Hz for 3000?ms, varying in magnitude 0 to 30?dB above threshold). At all frequencies from 16 to 125?Hz, absolute thresholds for the hand were significantly lower than those for the finger. Changes in threshold as a function of masker level were used to estimate the thresholds of three psychophysical channels (i.e. P, NP I, and NP II channels). Increased vibrotactile sensitivity of the hand compared to the finger seems to be not entirely due to increased spatial summation via the Pacinian system (P channel); non-Pacinian system (NP I and NP II channels) also contributed to perception. Differing transmission of vibration between the hand and the finger may have also influenced the thresholds.     

Independent responses of Pacinian and Non-Pacinian systems with hand-transmitted vibration detected from masked thresholds

This study was designed to identify psychophysical channels responsible for the detection of hand-transmitted vibration. Perception thresholds for vibration (16, 31.5, 63 and 125 Hz sinusoidal for 600 ms) at the distal phalanx of the middle finger and the whole hand were determined with and without simultaneous masking stimuli (1/3 octave bandwidth Gaussian random vibration centered on either 16 Hz or 125 Hz for 3000 ms, varying in magnitude 0 to 30 dB above threshold). At all frequencies from 16 to 125 Hz, absolute thresholds for the hand were significantly lower than those for the finger. Changes in threshold as a function of masker level were used to estimate the thresholds of three psychophysical channels (i.e. P, NP I, and NP II channels). Increased vibrotactile sensitivity of the hand compared to the finger seems to be not entirely due to increased spatial summation via the Pacinian system (P channel); non-Pacinian system (NP I and NP II channels) also contributed to perception. Differing transmission of vibration between the hand and the finger may have also influenced the thresholds.     

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.     

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.     

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.     

Vibrotactile thresholds at the fingertip, volar forearm, large toe, and heel

Thresholds for the perception of vibration vary with location on the body due to the organization of tactile channels in hairy and non-hairy skin, and variations in receptor density. This study determined vibration thresholds at four locations on the body with two different contactors so as to assist the identification of the tactile channel determining the threshold at each location. Vibrotactile thresholds at six frequencies from 8 to 250 Hz were measured on the distal phalanx of the index finger, the volar forearm, the large toe, and the heel with two contactors: (i) a 1-mm diameter circular probe with a 1-mm gap to a fixed circular surround (i.e., 7.1-mm(2) excitation area), and (ii) a 6-mm diameter circular probe with a 2-mm gap to a fixed circular surround (i.e., 79-mm(2) excitation area). At all frequencies and with both contactors, thresholds on the fingertip were lower than thresholds on the volar forearm, the large toe, and the heel, consistent with a greater density of mechanoreceptors at the fingertip. Thresholds with the larger contactor were lower than thresholds with the smaller contactor on the fingertip at high frequencies (63, 125, and 250 Hz), on the large toe (except at 250 Hz), on the heel (at all frequencies), and on the volar forearm at 250 Hz. It is concluded that at least two tactile channels (Pacinian from 63 to 250 Hz, and non-Pacinian from 8 to 31.5 Hz) determined vibrotactile thresholds at the fingertip, whereas non-Pacinian channels had a dominant influence on vibrotactile thresholds at the volar forearm. The role of Pacinian and non-Pacinian channels could not be confirmed at the large toe or the heel despite some evidence of spatial summation.     

Time course and action spectrum of vibrotactile adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation. In Experiment 2, test stimuli of 10 Hz and 50 Hz were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min. In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

Time Course and Action Spectrum of Vibrotactile Adaptation

In a series of experiments designed to explore the processes underlying adaptation of the sense of flutter-vibration, vibrotactile threshold was measured on the pad of the index finger, using Békésy tracking. Unadapted thresholds were first measured, for a number of frequencies (4-90 Hz) and contactor sizes (1-8 mm diameter). As expected, these measurements indicated the presence of (1) a Pacinian system possessing spatial summation and increasing in sensitivity, as frequency was raised, at the rate of 12 dB/octave; and (2) a non-Pacinian system showing little spatial summation, and with a frequency characteristic matching that of the NP I mechanism of Bolanowski et al. (1988). These baseline data of Experiment 1 guided the selection of stimulus parameters for subsequent experiments, in which threshold for a test stimulus was measured before, during, and after periods of vibrotactile adaptation.

In Experiment 2, test stimuli of 10 H_z and 50 H_z were combined factorially with 30-dB SL adapting stimuli of the same two frequencies. When the test stimulus was 10 Hz, the two adapting frequencies were equally effective in raising threshold; however, when the 50-Hz test stimulus was used, the 50-Hz adapting stimulus raised threshold by a greater amount than did the 10-Hz adapter. These results confirm on the finger the independence of adaptation in Pacinian and non-Pacinian channels, a result previously established on the thenar by other workers. For all four frequency combinations, threshold rose exponentially with a time constant of 1.5-2 min.

In Experiment 3, an action spectrum was determined, showing the adapting amplitude needed at each of a series of frequencies to raise the threshold of a 10-Hz stimulus by 10 dB; this spectrum was essentially flat from 30 to 90 Hz. The results, taken in conjunction with what is known about rapidly adapting cutaneous mechanoreceptors, imply that the effectiveness of an adapting stimulus is not determined solely by the amount of activity it generates in first-order afferents.     

Pitch detection of dynamic iterated rippled noise by humans and a modified auditory model

Iterated ripple noise (IRN) is a broadband noise with temporal regularities, which can give rise to a perceptible pitch. Since the perceptual pitch to noise ratio of these stimuli can be altered without substantially altering their spectral content, they have been useful in exploring the role of temporal processing in pitch perception [Yost, W.A., 1996. Pitch strength of iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3329-3335; Patterson, R.D., Handel, S.,Yost, W.A., Datta, A.J., 1996. The relative strength of the tone and noise components in iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3286-3294]. A generalised IRN algorithm is presented, in which multiple time varying temporal correlations can be defined. The resulting time varying pitches are perceptually very salient. It is also possible to segregate and track multiple simultaneous time varying pitches in these stimuli. Temporal auditory models have previously been shown to account for the perception of IRNs with static delays [Patterson, R.D., Handel, S.,Yost, W.A., Datta, A.J., 1996. The relative strength of the tone and noise components in iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3286-3294]. Here we show that some simple modifications to one such model [Meddis R., Hewitt, M.J., 1991. Virtual pitch and phase sensitivity of a computer model of the auditory periphery I. Pitch identification, J. Acoust. Soc. Am. 89, 2866-2882] allow it to track moving correlations, and also improve its performance in response to static correlations.     

Vibrotactile difference thresholds: Effects of vibration frequency,vibration magnitude,contact area,and body location

It has not been established whether the smallest perceptible change in the intensity of vibrotactile stimuli depends on the somatosensory channel mediating the sensation. This study investigated intensity difference thresholds for vibration using contact conditions (different frequencies, magnitudes, contact areas, body locations) selected so that perception would be mediated by more than one psychophysical channel. It was hypothesized that difference thresholds mediated by the non-Pacinian I (NPI) channel and the Pacinian (P) channel would differ. Using two different contactors (1-mm diameter contactor with 1-mm gap to a fixed surround; 10-mm diameter contactor with 2-mm gap to the surround) vibration was applied to the thenar eminence and the volar forearm at two frequencies (10 and 125?Hz). The up-down-transformed-response method with a three-down-one-up rule provided absolute thresholds and also difference thresholds at various levels above the absolute thresholds of 12 subjects (i.e., sensation levels, SLs) selected to activate preferentially either single channels or multiple channels. Median difference thresholds varied from 0.20 (thenar eminence with 125-Hz vibration at 10?dB SL) to 0.58 (thenar eminence with 10-Hz vibration at 20?dB SL). Median difference thresholds tended to be lower for the P channel than the NPI channel. The NPII channel may have reduced difference thresholds with the smaller contactor at 125?Hz. It is concluded that there are large and systematic variations in difference thresholds associated with the frequency, the magnitude, the area of contact, and the location of contact with vibrotactile stimuli that cannot be explained without increased understanding of the perception of supra-threshold vibrotactile stimuli.     

Vibrotactile thresholds at the sole of the foot: effect of vibration frequency and contact location

Studies of vibration perception in the glabrous skin of the human hand have identified four mechanoreceptor channels, with each channel showing characteristic variations in thresholds with variations in the frequency of vibration and the area of vibration excitation. To advance understanding of the channels mediating vibration perception on the sole of the foot, this study determined how thresholds depend on the frequency of vibration, the location on the foot (the big toe, the ball of the foot, and the heel), and the gap between a vibrating probe and a fixed surround. Thresholds at the three locations were obtained at the 12 preferred one-third octave centre frequencies from 20 to 250 Hz using a 6-mm diameter probe with both a 10-mm and a 20-mm diameter surround. With the 10-mm surround, the displacement thresholds at all three locations showed flat responses from 20 to 40 Hz. With both the 10-mm and the 20-mm surround, the displacement thresholds at the three locations showed "U-shaped" responses from 40 to 250 Hz. Relative to thresholds obtained with the 20-mm surround, thresholds obtained with the 10-mm surround were lower at the toe and the heel with 20- and 25-Hz vibration, but higher at the ball of the foot with 31.5- to 250-Hz vibration. It is concluded that absolute thresholds for the perception of vibration at the sole of the foot show important variations with location and with contact conditions and tend to be mediated by the NP I channel in the range from about 20 to 40 Hz and the P channel from about 40 to 250 Hz.     

Vibrotactile thresholds at the sole of the foot: Effect of vibration frequency and contact location

Studies of vibration perception in the glabrous skin of the human hand have identified four mechanoreceptor channels, with each channel showing characteristic variations in thresholds with variations in the frequency of vibration and the area of vibration excitation. To advance understanding of the channels mediating vibration perception on the sole of the foot, this study determined how thresholds depend on the frequency of vibration, the location on the foot (the big toe, the ball of the foot, and the heel), and the gap between a vibrating probe and a fixed surround. Thresholds at the three locations were obtained at the 12 preferred one-third octave centre frequencies from 20 to 250?Hz using a 6-mm diameter probe with both a 10-mm and a 20-mm diameter surround. With the 10-mm surround, the displacement thresholds at all three locations showed flat responses from 20 to 40?Hz. With both the 10-mm and the 20-mm surround, the displacement thresholds at the three locations showed “U-shaped” responses from 40 to 250?Hz. Relative to thresholds obtained with the 20-mm surround, thresholds obtained with the 10-mm surround were lower at the toe and the heel with 20- and 25-Hz vibration, but higher at the ball of the foot with 31.5- to 250-Hz vibration. It is concluded that absolute thresholds for the perception of vibration at the sole of the foot show important variations with location and with contact conditions and tend to be mediated by the NP I channel in the range from about 20 to 40?Hz and the P channel from about 40 to 250?Hz.     

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 (15d`, 30d`, and 40d`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 P_h (Pacinian, hairy skin), NP_{h low} (non-Pacinian, hairy skin, low frequencies) and NP_{h 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.     

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.     

Time-dependence of SI RA neuron response to cutaneous flutter stimulation

Spike discharge activity of RA-type SI cortical neurons was recorded extracellularly in anesthetized monkeys and cats. Multiple applications (trials) of 10-50 Hz sinusoidal vertical skin displacement stimulation ("flutter") were delivered to the receptive field (RF). Analysis revealed large and systematic temporal trends not only in SI RA neuron responsivity (measured as spikes/s and as spikes/stimulus cycle), but also in entrainment, and in phase angle of the entrained responses. In contrast to SI RA neurons, the response of RA skin afferents to comparable conditions of skin flutter stimulation exhibited little or no dynamics. The occurrence and form of the SI RA neuron response dynamics that accompany skin flutter stimulation are shown to depend on factors such as stimulus frequency and the locus of the recording site in the global cortical response pattern. Comparison of recordings obtained in near-radial vs tangential microelectrode penetrations further reveals that the SI RA neuron response dynamics that occur during skin flutter stimulation are relatively consistent within, but heterogeneous across column-sized regions. The observed SI RA neuron response dynamics are suggested to account, in part, for the improved capacity to discriminate stimulus frequency after an exposure ("adaptation") to skin flutter stimulation (Goble and Hollins, J Acoust Soc Am 96: 771-780, 1994). Parallels with recent proposals about the contributions to visual perception of short-term primary sensory cortical neuron dynamics and synchrony in multineuron spike activity patterns are identified and discussed.