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.     

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.     

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.     

Reductions in finger blood flow in men and women induced by 125-Hz vibration: association with vibration perception thresholds

Vibration of one hand reduces blood flow in the exposed hand and in the contralateral hand not exposed to vibration, but the mechanisms involved are not understood. This study investigated whether vibration-induced reductions in finger blood flow are associated with vibrotactile perception thresholds mediated by the Pacinian channel and considered sex differences in both vibration thresholds and vibration-induced changes in digital circulation. With force and vibration applied to the thenar eminence of the right hand, finger blood flow and finger skin temperature were measured in the middle fingers of both hands at 30-s intervals during seven successive 4-min periods: 1) pre-exposure with no force or vibration, 2) pre-exposure with force, 3) vibration 1, 4) rest with force, 5) vibration 2, 6) postexposure with force, and 7) recovery with no force or vibration. A 2-N force was applied during periods 2-6 and 125-Hz vibration at 0.5 and 1.5 ms(-2) root mean square (r.m.s.; unweighted) was applied during periods 3 and 5, respectively. Vibrotactile thresholds were measured at the thenar eminence of right hand using the same force, contact conditions, and vibration frequency. When the vibration magnitude was greater than individual vibration thresholds, changes in finger blood flow were correlated with thresholds (with both 0.5 and 1.5 ms(-2) r.m.s. vibration): subjects with lower thresholds showed greater reductions in finger blood flow. Women had lower vibrotactile thresholds and showed greater vibration-induced reductions in finger blood flow. It is concluded that mechanoreceptors responsible for mediating vibration perception are involved in the vascular response to vibration.     

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.     

Effects of temperature on the subjective magnitude of vibration

Absolute magnitude estimation (AME) was used to determine the effects of skin temperature on the subjective magnitude of vibration delivered to the thenar eminence of the right hand. Measurements were made at three frequencies chosen to selectively activate cutaneous mechanoreceptor channels, namely NP I and NP III (Meissner and Merkel cell-neurite receptors, respectively) at 15 Hz, Pacinian (250 Hz, 400 Hz) and NP III (Ruffini endings) at 400 Hz. Skin temperatures at 15, 20 and 40 degrees C were tested at 11 suprathreshold displacement levels. It was concluded that the subjective magnitude of vibration is influenced by temperature in the Pacinian channel, but in the NP I and NP III channels the temperature of the skin did not have an effect upon judgements of subjective magnitude. This is consistent with earlier work by Verrillo and Bolanowski (J Acoust Soc Am 80: 528-532, 1986) and Bolanowski et al. (J Acoust Soc Am 84: 1680-1694, 1988); both studies show only modest effects of temperature at threshold at frequencies below approximately 40 Hz.     

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.     

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.     

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.     

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.     

Vibrotactile thresholds of the Non-Pacinian I channel: I. Methodological issues

Thresholds of the Non-Pacinian I (NP I) channel were measured using a two-interval forced-choice paradigm, a technique independent of the subject's criterion. The studies were performed using the terminal phalanx of the human middle finger with a 40-Hz vibratory stimulus. Unlike most of the previous experiments performed in our laboratory, a contactor surround was not used. This was done to enable comparison with population models of mechanoreceptive fibers in the literature. Since the Pacinian (P) channel and NP I channel have similar vibrotactile thresholds at 40?Hz, a forward-masking procedure was used to elevate the thresholds of the P channel with respect to the NP I channel. While it has been established that the Pacinian fibers are entrained at high stimulus levels, the P channel can be perceptually masked using a 250-Hz stimulus presented prior to the 40-Hz test stimulus. The masking functions were found to be approximately linear on log-log axes and the threshold shifts were found to increase as the masking-stimulus levels increased. The results are discussed in relation to previous studies that were performed at various stimulation sites by using a contactor surround or not. A companion paper presents the variation of NP I-channel thresholds, measured using the methods described herein, and addresses the effects of stimulation along the proximo-distal axis of the phalanx. The companion paper also discusses the predictions of a computational model, recently proposed, in light of the empirical results presented.     

Vibrotactile thresholds of the Non-Pacinian I channel: I. Methodological issues

Thresholds of the Non-Pacinian I (NP I) channel were measured using a two-interval forced-choice paradigm, a technique independent of the subject's criterion. The studies were performed using the terminal phalanx of the human middle finger with a 40-Hz vibratory stimulus. Unlike most of the previous experiments performed in our laboratory, a contactor surround was not used. This was done to enable comparison with population models of mechanoreceptive fibers in the literature. Since the Pacinian (P) channel and NP I channel have similar vibrotactile thresholds at 40?Hz, a forward-masking procedure was used to elevate the thresholds of the P channel with respect to the NP I channel. While it has been established that the Pacinian fibers are entrained at high stimulus levels, the P channel can be perceptually masked using a 250-Hz stimulus presented prior to the 40-Hz test stimulus. The masking functions were found to be approximately linear on log-log axes and the threshold shifts were found to increase as the masking-stimulus levels increased. The results are discussed in relation to previous studies that were performed at various stimulation sites by using a contactor surround or not. A companion paper presents the variation of NP I-channel thresholds, measured using the methods described herein, and addresses the effects of stimulation along the proximo-distal axis of the phalanx. The companion paper also discusses the predictions of a computational model, recently proposed, in light of the empirical results presented.     

Effect of contact location on vibrotactile thresholds at the fingertip

Vibrotactile thresholds depend on the characteristics of the vibration, the location of contact with the skin, and the geometry of the contact with the skin. This experimental study investigated vibrotactile thresholds (from 8 to 250 Hz) at five locations on the distal phalanx of the finger with two contactors: (i) a 1-mm diameter circular probe (0.78-mm(2) area) 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 (28-mm(2) area) with a 2-mm gap to a fixed circular surround (i.e., 79-mm(2) excitation area). With both contactors, especially the smaller contactor at low frequencies (i.e., 8, 16, and 31.5 Hz), thresholds decreased towards the tip of the finger, although there was little variation around the whorl. With low frequencies of vibration, and at all five locations on the finger, similar thresholds were obtained with both contactors, consistent with the NPI channel not changing in sensitivity with a change in the area of stimulation. At high frequencies (i.e., 63, 125, and 250 Hz), thresholds were lower with the larger area of stimulation at all locations, except at the extreme tip of the finger, consistent with spatial summation in the Pacinian channel. It is concluded that with a 6-mm diameter contactor, moderate variations in location around the whorl have little influence on the measured thresholds. With the 1-mm diameter contactor there were greater variations in thresholds and extreme locations, near the nail and the distal interphalangeal joint, may be unsuitable for investigating sensorineural disorders.     

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.     

Frequency and site-dependent variations in vibration detection thresholds on the face

An adaptive psychophysical procedure was used to estimate the vibration detection threshold at seven spatially matched sites on the two sides of the face and at one scalp site. Repeated measurements over six testing sessions were made for stimuli vibrating at 1, 10 and 100 Hz for each of 21 neurologically healthy, young adult females. Approximately 14 stimulus trials were required to obtain each estimate of the threshold amplitude. Thresholds varied as a function of frequency ( p < 0.0001), side ( p < 0.001) and site ( p < 0.0001). Compared to stimulation at 100 Hz at which the estimates were lowest, thresholds were 3.1 times greater at 10 Hz and 5.4 times greater at 1 Hz. Thresholds were lowest on the vermilion and highest on the cheek and chin. The preauricular skin and scalp exhibited an intermediate level of sensitivity. Whereas thresholds were comparable on the two sides of the face for stimulation at 1 Hz, they averaged 1.33 times greater on the right side for stimulation at 10 and 100 Hz. Moreover, thresholds obtained during the last two sessions were 16% higher than those obtained during the first two sessions ( p < 0.02), suggesting that subjects on average became more conservative in reporting the presence of the stimulus. The sensitivity in discriminating differences in tactile function favors use of the rapidly administered testing procedure in a clinical setting.     

Frequency and site-dependent variations in vibration detection thresholds on the face

An adaptive psychophysical procedure was used to estimate the vibration detection threshold at seven spatially matched sites on the two sides of the face and at one scalp site. Repeated measurements over six testing sessions were made for stimuli vibrating at 1, 10 and 100 Hz for each of 21 neurologically healthy, young adult females. Approximately 14 stimulus trials were required to obtain each estimate of the threshold amplitude. Thresholds varied as a function of frequency (p < 0.0001), side (p < 0.001) and site (p < 0.0001). Compared to stimulation at 100 Hz at which the estimates were lowest, thresholds were 3.1 times greater at 10 Hz and 5.4 times greater at 1 Hz. Thresholds were lowest on the vermilion and highest on the cheek and chin. The preauricular skin and scalp exhibited an intermediate level of sensitivity. Whereas thresholds were comparable on the two sides of the face for stimulation at 1 Hz, they averaged 1.33 times greater on the right side for stimulation at 10 and 100Hz. Moreover, thresholds obtained during the last two sessions were 16% higher than those obtained during the first two sessions (p < 0.02), suggesting that subjects on average became more conservative in reporting the presence of the stimulus. The sensitivity in discriminating differences in tactile function favors use of the rapidly administered testing procedure in a clinical setting.     

Effect of contact location on vibrotactile thresholds at the fingertip

Vibrotactile thresholds depend on the characteristics of the vibration, the location of contact with the skin, and the geometry of the contact with the skin. This experimental study investigated vibrotactile thresholds (from 8 to 250?Hz) at five locations on the distal phalanx of the finger with two contactors: (i) a 1-mm diameter circular probe (0.78-mm² area) with a 1-mm gap to a fixed circular surround (i.e., 7.1-mm² excitation area), and (ii) a 6-mm diameter circular probe (28-mm² area) with a 2-mm gap to a fixed circular surround (i.e., 79-mm² excitation area). With both contactors, especially the smaller contactor at low frequencies (i.e., 8, 16, and 31.5?Hz), thresholds decreased towards the tip of the finger, although there was little variation around the whorl. With low frequencies of vibration, and at all five locations on the finger, similar thresholds were obtained with both contactors, consistent with the NPI channel not changing in sensitivity with a change in the area of stimulation. At high frequencies (i.e., 63, 125, and 250?Hz), thresholds were lower with the larger area of stimulation at all locations, except at the extreme tip of the finger, consistent with spatial summation in the Pacinian channel. It is concluded that with a 6-mm diameter contactor, moderate variations in location around the whorl have little influence on the measured thresholds. With the 1-mm diameter contactor there were greater variations in thresholds and extreme locations, near the nail and the distal interphalangeal joint, may be unsuitable for investigating sensorineural disorders.     

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.