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Pacinian corpuscles (PCs) in cat mesentery have been studied extensively to help determine the structural and functional bases of tactile mechanotransduction. Although we, like many other investigators, have found that the mesenteric receptors are anatomically very similar to those found in mammalian skin, few physiological characteristics of the mesenteric PCs and those of the skin have been compared. Action-potential rate-amplitude and frequency characteristics (10 Hz-1 KHz), as well as interval (IH) and peri-stimulus-time (PSTH) histograms in response to sinusoidal displacements were obtained from nerve fibers innervating mesenteric PCs and from PC fibers innervating cat glabrous skin. The intensity characteristics obtained on both preparations showed similar response profiles, including equal slopes for low stimulus intensities (approximately 10, with impulse ratios/20 dB displacement) and one and two impulse/cycle entrainment. The frequency characteristics of both groups were U-shaped with similar low-frequency slopes (-12.5 dB/octave) and bandwidths (Q(3dB) = 1.4). The best frequency for both the tactile PCs' and mesenteric PCs was 250 Hz, which is in the expected range. The IHs showed entrainment and the PSTHs showed neither transient responses nor adaptation to steady-state sinusoidal stimuli. The functional similarity between mesenteric PCs' nerve responses and those of tactile PC afferents, as well as the receptors' anatomical similarity, lead us to suggest that the mesenteric PC can act as a model for those in the skin. Furthermore, since the frequency characteristics of the two PC types are similar, it is concluded that the skin, while attenuating stimulus intensity, does not impart temporal filtering of vibratory stimuli.

The Merkel cell-neurite complex is considered to be one class of mechanoreceptors in the skin. Merkel cells are innervated by slowly adapting type I (SAI) tactile nerve fibers. In this paper, the detailed distribution of Merkel cells is studied by immunohistochemical labeling of the monkey (Macaca fascicularis) digital glabrous skin. Specific morphometric variables (density of intermediate epidermal ridges and Merkel cells, distance between skin surface and ridge tips and bases, maximum and average cell counts per ridge, distance between cells and ridges) were measured by a combination of light/fluorescence microscopy and computer-image analysis. The morphometric results were similar for each digit of the monkey's hand. Next, the anatomical data were used to form a three-dimensional reconstruction of the Merkel-cell distribution in the fingertip skin. A patch of the distal-pad surface was then computationally flattened to obtain the two-dimensional distribution of Merkel cells. Based on previous anatomical and physiological data, SAI fibers were simulated to innervate clusters of Merkel cells in the distal-pad surface. On average, 28 cells were innervated by a single fiber. The resulting anatomical model may be used to estimate the population response of SAI fibers by incorporating spike generation.

The role of the capsule encasing the Pacinian corpuscle's (PC's) neurite, where mechanotransduction occurs, may be more than mechanical. The inner core of the PC's capsule consists of lamellar cells that are of Schwann-cell origin. Previously, we found both voltage-gated Na+ and K+ channels in these inner-core lamellae. Research on astrocytes and Schwann cells shows bidirectional signaling between glia and neurons, a major component of which is glutamate. Furthermore, Merkel cells show positive immunoreactivity for glutamate receptor mGluR5, and the glutamate-receptor antagonist kynurenate greatly decreases the static activity of the slowly adapting neurons of Merkel cell-neurite complexes. To investigate the possibility of glutaminergic interaction in PCs, we applied antibodies to glutamate, glutamate receptors, glutamate transporters, and SNARE proteins to cat mesenteric PC sections. Positive labeling was seen in the inner-core lamellae, at inter-lamellar connections, where the lamellae contact the membrane of the neurite and at the lamellar tips. The presence of these proteins on the lamellae and neurite membranes, demonstrated both with immunofluorescent light microscopy as well as immunogold electron microscopy, suggests a chemical, possibly bidirectional, interaction between the lamellar cells and the neurite. Thus, the capsule of the PC, apart from having a mechanical filtering function, may also provide an environment for lamellar-neurite interaction, perhaps acting as a neuro-modulator of the initiation, and/or continuation, of the mechanical-electrical transduction process. At the very least, the presence of the aforementioned proteins suggest some sort of "synaptic-like" activity in these mechanoreceptors, which up until now has not been considered possible.

Static displacements in Pacinian corpuscles (PCs) were measured using video microscopy. Mechanical stimuli of 10-40 microm steps were applied to the PC capsule surfaces using cylindrical contacts with different diameters. Displacements parallel to the stimulation axis were measured at various locations in the focal plane of the optical setup. In contrast to previous data in the literature, the displacements within the corpuscle were found to be linearly related to the indentation amplitude. Displacements decreased as a function of lamella depth, with a more negative slope close to the surface and less negative slope at deeper locations. The experimental data were compared to the predictions of a previous mechanical model, and to the results of two new models: (1) elastic semi-infinite continuum model; (2) ovoid isotropic finite-element model. Although the previous model did not specify displacement boundary conditions, it predicted the current experimental results well. On the other hand, the experimental displacements were found to be smaller than those predicted by the semi-infinite continuum and finite-element models. However, both semi-infinite continuum and finite-element models yielded close results, which show that the three-dimensional ovoid geometry of the corpuscle is not the primary factor for determining the displacements in physiological conditions. Furthermore, simulations with the finite-element model using a wide range of material properties yielded similar results. This supports the hypothesis that a homogeneous isotropic model for the PC cannot predict experimental results. The modeling analyses suggest that the experimental results are largely affected by the displacement of the incompressible interlamellar fluid and the layered structure of the corpuscle.

Strength loss following disuse may result from alterations in muscle and/or neurological properties. In this paper, we report our findings on human plantar flexor neurological properties following 4 wk of limb suspension [unilateral lower limb suspension (ULLS)], along with the effect of motor imagery (MI) training on these properties. In the companion paper (Part I), we report our findings on the changes in skeletal muscle properties. Additionally, in the present paper, we analyze our findings to determine the relative contribution of neural and muscular factors in strength loss. Measurements of central activation, the H-reflex, and nerve conduction were made before and after 4 wk of ULLS (n = 18; 19-28 yr). A subset of the subjects (n = 6) performed PF MI training 4 days/wk. Following ULLS, we observed a significant increase in the soleus H-reflex (45.4 +/- 4.0 to 51.9 +/- 3.7% expressed relative to the maximal muscle action potential). Additionally, there were longer intervals between the delivery of an electrical stimulus to the tibial nerve and the corresponding muscle action potential (M-wave latency; mean prolongation 0.49 ms) and H-reflex wave (H-wave latency; mean prolongation 0.46 ms). The efficacy of MI on strength was ambiguous, with no significant effect detected (although a modest effect size was observed; eta2 = 0.18). These findings suggest that unweighting induces plastic changes in neural function that appear to be spatially distributed throughout the nervous system. In terms of the relative contribution of neural and muscular factors regulating strength loss, we observed that neural factors (primarily deficits in central activation) explained 48% of the variability in strength loss, whereas muscular factors (primarily sarcolemma function) explained 39% of the variability.

A computational model based on previous physiological and psychophysical data is presented for the human Pacinian (P) psychophysical channel. The model can predict the probability of detection in simple psychophysical tasks, and hence psychometric functions and thresholds. The model simulates stimulating variable and fixed glabrous skin sites with different-sized contactors and includes spatial variation of monkey P-fiber sensitivities. Therefore, it is especially suitable for studying spatial summation, i.e. the improvement of threshold with increasing contactor area. Selective contributions of neural integration (n.i.) and probability summation (p.s.) are also incorporated into the model. Model predictions are compared to psychophysical results of Gescheider et al. (2005). The performance of the model regarding the effects of contactor size is very good. In addition to predicting approximately 3 dB improvement of thresholds when the contactor area is doubled, the model also reveals nonlinear contributions of p.s. and n.i. Furthermore, the model asserts that thresholds are largely governed by neural integration when small contactors are used. These and other findings discussed in the article show that the presented model is a helpful tool for formulating testable hypotheses. Although the model can also simulate some temporal summation effects, simulation results do not conform well to previous data on temporal response properties. Thus, the model needs to be refined in that respect.

The interaction of excitation and inhibition in responses due to attentional mechanisms in the visual system has been investigated. The studies reported herein use the tactile system of humans to test a specific hypothesis about the processes of attention that have never been directly addressed. Both exogenous and endogenous Inhibition of Return (IOR) reaction-time paradigms with a 100 Hz, 35 microm of peak displacement amplitude were used. In these experiments multiple Stimulus Onset Asynchronies were tested which made it difficult for subjects to learn timing patterns. We tested whether a detection time to a target which is to be attended to is a composite of at least two underlying mechanisms. These mechanisms were explored using exogenous and endogenous IOR experiments. It is hypothesized that these mechanisms work in a push-pull fashion: one deploying attention when new events occur, and the other withdrawing attention ("Disengagement") after it has been deployed. Based on the results, a new hypothesis is proposed stating that one form of attention (selective attention) competes with IOR in determining the time taken to detect a target in the tactile system.

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.

The firing-rate-based population model for rapidly-adapting (RA) mechanoreceptive fibers by Güçlü and Bolanowski is extended by including temporal-response properties of RA fibers. This representation allows for the generation of action-potential (spike) times for each fiber when a sinusoidal, steady-state stimulus is applied onto the skin. Signal detection theory was used to predict human psychophysical thresholds. Specifically, the effects of sensorineural innervation pattern, stimulus-contactor location and selected decision rules on the model predictions were studied. The predicted thresholds were lowest when the decision rule was one spike and highest when many active fibers were required for detection. These predictions were empirically tested by measuring vibrotactile thresholds of the Non-Pacinian I (NP I) channel, which required the special techniques discussed in the preceding article. Although the model predicted thresholds to decrease distally due to the known innervation density which is higher distally, the thresholds of the NP I psychophysical channel were found to be approximately constant (20-25 dB re 1 microm peak amplitude) from the proximal site on the terminal phalanx to the most distal portion. Interestingly, the mechanical impedance of the skin was found not to be constant along the proximo-distal axis. This latter result implies that the space-invariant mechanical attenuation function used in the model may not be valid at every location on the fingertip. Because of this, the discrepancy between the model's predictions and the psychophysical results may be reconciled.

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.

Common confounding factors for polarimetric concentration measurements include additional optical rotations from unknown optically active molecules, linear birefringence of the medium, and path length variability. We show that by approximating Drude's equation and taking several measurements from the same sample at different wavelengths, the error due to confounding rotations in the measurements can theoretically be canceled. The analysis is developed with regard to glucose sensing in aqueous humor. First, we show that the optical rotatory dispersions of the known molecules in bovine aqueous humor could be represented by Drude's equations. Then, the total optical rotation is approximated by a function combining Drude's equations for the major contributors in the sample, i.e., glucose, glutamine, fructose, and phenylalanine. The concentration-related unknown coefficients in the approximating function are found by constrained nonlinear optimization of the function at different wavelengths. This technique is tested on a published data set and four alterations of those data: (1) concentrations randomly varied within narrow limits, (2) similar to alteration 1 but with significantly elevated glucose concentration, (3) similar to alteration 1 but with significantly decreased glucose concentration, and (4) concentrations randomly varied within wider limits than alteration 1. The method produces very accurate glucose-concentration estimates in all of these data sets. The relative error was smaller than 1% in all except the low-glucose sample (1.4%). This method may prove useful in noninvasive glucose measurement in humans., ((c) 2004 Society of Photo-Optical Instrumentation Engineers.)

Rapidly-adapting (RA) mechanoreceptive fibers, which are associated with Meissner corpuscles, mediate one component of the neural information that contributes to the sense of touch. Responses of cat RA fibers subject to 40-Hz sinusoidal stimulation were modeled as a Markov process. Since an RA fiber generates one, two or no spikes in each cycle of the stimulus, the fiber's activity was considered to exist in one of these three possible states. By analyzing empirically generated spike trains, the probability of each state and the probabilities of transitions between the three states were found as a function of the average firing rate of the fiber. The average firing rate depends on the stimulus amplitude. In addition, the phase of each spike with respect to the stimulus cycle was represented by a Laplace distribution. Based on empirical data, the mean and the standard deviation of this distribution decrease as the stimulus amplitude is increased. The entire stochastic model was implemented on a computer to simulate the responses of RA fibers. The post-stimulus time, inter-spike interval and period histograms generated from the simulations match the histograms obtained from the empirical data well as quantified by relative errors. This temporal model can be combined with a population model for average rate to derive a spatio-temporal description of the responses of somatosensory afferents. The effects of changing the stimulation frequency are discussed.

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.

A series of experiments are described in which magnitude estimates of the perceived size of steel balls were made when the balls were actively rolled between the fingertip and several other body sites (thumb, thenar eminence, forearm). This movement, called scripting, involves actively moving an object by a touching surface over another surface of the body which is passively being touched. We define this active/passive activity as "intra-active touch" and the results show that the perceptual size of the balls is dependent upon the body part passively being activated. An additional series of experiments decoupled the actively generated and passively received tactile information by having subjects either perform the scripting on another individual's body site or by having the other individual roll the balls on the subject's various sites. The latter experiments showed that the passive body can contribute to the overall impression of the size of the balls, but only when the intra-active touching involved the glabrous skin of the hands. Intra-active touch between the active finger and the passively touched hairy skin of the forearm showed no effect of the touched surface on the perceived size of the balls. The results suggest that the mechanisms of intra-active touch are different when glabrous skin activates glabrous skin than when glabrous skin activates hairy skin.

The goal of this study is to establish a link between somatosensory physiology and psychophysics at the probabilistic level. The model for a population of monkey rapidly adapting (RA) mechanoreceptive fibers by Güçlü and Bolanowski (2002) was used to study the probability of stimulus detection when a 40 Hz sinusoidal stimulation is applied with a constant contactor size (2 mm radius) on the terminal phalanx. In the model, the detection was assumed to be mediated by one or more active fibers. Two hypothetical receptive field organizations (uniformly random and gaussian) with varying average innervation densities were considered. At a given stimulus-contactor location, changing the stimulus amplitude generates sigmoid probability-of-detection curves for both receptive field organizations. The psychophysical results superimposed on these probability curves suggest that 5 to 10 active fibers may be required for detection. The effects of the contactor location on the probability of detection reflect the pattern of innervation in the model. However, the psychophysical data do not match with the predictions from the populations with uniform or gaussian distributed receptive field centers. This result may be due to some unknown mechanical factors along the terminal phalanx, or simply because a different receptive field organization is present. It has been reported that human observers can detect one single spike in an RA fiber. By considering the probability of stimulus detection across subjects and RA populations, this article proves that more than one active fiber is indeed required for detection.

A novel hierarchical clustering algorithm was applied to the distribution of Meissner corpuscles in the skin of mammals. This method, called end-to-end linkage (EEL), is useful for grouping data that consists of chain-like contingencies in the multivariable space. Unlike the traditional techniques which uncover hyperspherical clusters (e.g. single linkage), EEL considers the shortest distance between the predefined end pairs of the two clusters as an inter-group distance. This scheme allows characterizing the internal structure of data better than other hierarchical techniques. The anatomical data used in the case study is important for studying the sense of touch. The results show a substantial improvement over the traditional single-linkage method. On average, the number of correctly classified corpuscles is increased to twice the number identified by the single-linkage method. EEL can also be used for analyzing other sensory modalities where geometric relationships need to be explored. In addition, the report contains corpuscle density and epidermal-ridge width data obtained from several species.

Experiments were performed in which the subjective size of steel balls was judged by the method of absolute magnitude estimation (AME). The balls were rolled actively ("scripting") by the right index finger pad of the subject over the passively receiving areas of the face, namely the forehead, cheek and the vermilion border of the lower lip. These areas were stimulated on the subject's own person (intra-active touch), on another person's face (interactive touch) and by another person upon the face of the subject (interactive touch). Results indicate that size perception appears to be governed primarily by the mechanical characteristics of the stimuli, including the balls and the sites on the face. When the subject's finger was used to script upon another's face, there were significant differences among or between the sites. When another person scripted upon the subject's face there were no significant differences among any of the sites. The results are discussed considering mode of stimulation and tissue mechanics.

Reconstructing neural-population responses in the form of spatial event plots assumes that the receptors are organized in a dense linear array. We have found that this assumption is not valid by determining the spatial organization of Meissner corpuscles (MCs) in the glabrous skin of both cat and monkey. The tissue was excised from animals that had been cardiac perfused with 4% paraformaldehyde. One-micrometer plastic sections revealed that the morphology of these receptors is different in the two animal species. However, in both species, they reside in approximately the same place in the dermal pegs of the skin, between the epidermal ridges, and electrophysiologically they both respond to ramp-and-hold stimuli with a rapidly adapting firing pattern. Thus, in this study we will refer to the receptors of the cat as "Meissner-like". In monkey, MCs are located in the dermal papillae between the epidermal limiting and intermediate ridges, forming orderly rows, the contours of which follow the overlying fingerprint. Although the average density of MCs is 45/mm2, they are distributed along the dermal pegs in such a manner as to give rise to three significantly ( p < 0.017) different average distances between corpuscles. We note that "fingerprints" vary in topography across the hand and this is also reflected in the underlying MC arrays. In the cat, these "Meissner-like" receptors display no specific pattern and have a density much lower than in the monkey. Cat glabrous skin does not have "fingerprints". The results emphasize that the spatial organization of tactile receptors must be taken into account when interpreting reconstructed population responses.

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 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.

Modeling population responses of nerve fibers requires statistical characterization of fiber-response properties. The rate/intensity characteristics of cat rapidly adapting (RA) fibers were fitted by four-parameter, piece-wise linear functions using nonlinear regression (n = 14; R2 > 0.958). The parameters were tested against the null hypothesis that they are log normally distributed. The test fail to reject this hypothesis (Kolmogorov-Smirnov p>0.477). However, a significant statistical difference was found between the specific lognormal distributions obtained from monkey (Johnson, J Neurophysiol 37: 48-72, 1974) and cat for all four parameters (Kolmogorov-Smirnov, p<0.0075, p<0.05, p<0.0001, p<0.00007). Although the stimulus contactor size was not the same in monkey and cat studies, the differences between monkey and cat fibers are attributed to anatomical differences in the glabrous sin of both species. Modeling studies suggest that the absolute firing thresholds of RA fibers have a right-skewed distribution because of the anatomical constraints present in both species' skin. Meissner corpuscles, which are the sensory end-organs of RA fibers, are likely to be found deeper in the skin within dermal papilla, therefore, the thresholds can be elevated. However, the thresholds are bounded at lower end, probably due to the epidermal junction that acts as a superficial mechanical barrier for these corpuscles.

The frequency responses of 11 rapidly adapting (RA) fibers in cat were studied by representing the average firing rate as a function of sinusoidal stimulus amplitude and stimulus frequency. Specifically, rate-intensity functions at different stimulation frequencies were fitted by four-parameter (a0, a1, a2, a3), piece-wise linear functions using nonlinear regression (n = 59; R2 > 0.877). Rate-intensity functions at intermediate frequencies were found by linear interpolation. The result of this analysis is rate-amplitude-frequency functions plotted as two-dimensional surfaces. The surfaces consist of five regions separated and sufficiently defined by four space curves. At 14 different frequencies, the statistical distribution of each rate-intensity-function parameter could be approximated by a particular lognormal distribution (n = 56; R2 > 0.796). The Kolmogorov-Smirnov test fails to reject this hypothesis for each combination of frequency and parameter (56 tests; p > 0.39). Therefore, at a given frequency, the variation of the parameters can be represented by lognormal distributions with specific means and standard deviations. Responses of six RA fibers, which are different from the data-set used for modeling, were compared with the stochastic model at different frequencies. The parameters of those fibers were tested against the null hypotheses that they were sampled from the particular parameter distributions dictated by the model. The Kolmogorov-Smirnov test fails to reject all the hypotheses at the alpha = 0.05 level (44 tests). At the alpha = 0.10 level, only a few test parameters were found to be departing from the model (a0 and a1 at 5 Hz; a2 at 20 Hz; a2 and a3 at 50 Hz). The remaining test parameters could be accurately described by the model. Having confirmed the validity of the model, the logarithmic means and the logarithmic standard deviations of the lognormally distributed rate-intensity-function parameters were estimated in the frequency range of 4-200 Hz. The rate-amplitude-frequency surfaces sampled from the established stochastic model completely characterize the rate responses of RA fibers to sinusoidal stimuli and are superior to tuning curves which require selecting criterion responses. The current rate-response model is promising for future computational work, especially on population modeling.

The experimental setup for generating spatial event plots (SEPs) from single mechanoreceptive fibers of the skin was computationally simulated. The generic fibers used in the simulations were similar to the rapidly adapting fibers (RAs), and had variable refractoriness and receptive-field size. The speed, lateral shift, and the contact width of the drum scanned across the receptive field of the fiber are adjustable parameters. The stimulus patterns used on the drum mimicked stimuli used by several other investigators. These were dot patterns, grating patterns, and the letter "E". First, the effects of simulation parameters on the SEPs were studied. The simulation output confirms the results of physiological experiments that SEPs contain information on the spatiotemporal resolution of the fiber. The next series of simulations involved generating SEPs of fibers obtained from the same or varying spatial distributions of receptive fields. Three hypothetical distributions were used: homogeneous rectangular, uniformly random, and Gaussian. The momentary population response at each case was found using the technique by Johansson and Vallbo (Brain Res 184: 353-366, 1980). The population responses were not isomorphic images of the stimulus patterns due to the variations in field sizes and locations. However, every fiber, no matter which distribution it came from, generated almost identical SEPs given similar response properties. Furthermore, the SEPs looked like the outline of the stimulus. These observations show that SEPs do not contain information about the population response. Therefore, reconstructing the population response using SEPs can result in misleading conclusions on central-nervous-system processing and should be viewed cautiously when formulating psychophysical/physiological linking hypotheses.

The population response of rapidly-adapting (RA) fibers is one component of the physiological substrate of the sense of touch. Herein, we describe a computational scheme based on the population-response model by K.O. Johnson (J. Neurophysiol. 37: 48-72, 1974) which we extended by permitting the capability to include the spatial distributions of receptors in the glabrous skin linked to RA fibers. The hypothetical cases simulated were rectangular, uniformly random and proximo-distally Gaussian distributions. Each spatial organization produced qualitatively distinct population-response profiles that also varied due to stimulus parameters. The effects of stimulus amplitude, average innervation density and contactor-probe location were studied by considering various response measures: number of active fibers, summated firing rate and the average firing rate of a subset of the modeled population. The outcome of the measures were statistically compared among simulated anatomical distributions. The response is the same for rectangular and uniformly random distributions, both of which have a homogeneous innervation density. However, the Gaussian distribution produced statistically different responses when the measure was not averaged over the subset population which represented the receptive field of a higher-order neuron. These results indicate that, as well as stimulus parameters, the anatomical organization is a significant determinant of the population response. Therefore, reconstructing population activity for testing psychophysical hypotheses must presently be done with care until the organization of the receptors within the skin has been clarified.

It has long been accepted that action potentials arising from Pacinian corpuscles (PCs) originate at the first node of Ranvier located within the PC and that the mechanotransduction events (receptor potentials) are formed by stretch-activated channels selectively sensitive predominantly to Na+. Also, it has been shown previously that tetrodotoxin (TTX) affects the receptor potential suggesting that transduction may involve voltage-sensitive Na+ channels. To determine whether voltage-sensitive Na+ channels exist in the membrane thought to be responsible for transduction, immunocytochemical studies were performed using polyclonal antibodies raised in rabbit against the alpha subunit of rat type I and type II voltage-gated sodium channels. The results show the presence of label on the neurite and axolemma, as well as in the node regions. Interestingly, labeling is also found on the inner and outer lamellae that form the non-neural accessory structure surrounding the neurite. The presence of this label in the surrounding lamellae suggests that voltage-sensitive Na+ channels, that are involved in both transduction and action-potential generation, may be made available to the neurite via transport from the lamellae, a mechanism perhaps operating in parallel to axoplasmic transport.

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.

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.

The effects of heat-induced pain on absolute thresholds, sensation magnitudes and amplitude-difference thresholds were measured at 10 and 100 Hz. Consistent with previous results, heat-induced pain elevated the absolute thresholds by approximately 8.0 dB and lessened the magnitudes of tactile sensations during pain as compared to the non-painful condition. In contrast to these effects, the discriminability of change in the intensity of the vibrotactile stimuli was unaffected by the presence of pain indicating that the effect of pain on tactile sensations is more likely due to sensory rather than cognitive processes (i.e., attention) and that the mechanisms underlying tactile sensitivity as compared to discriminability are different.

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 influence of heat- and cold-induced pain on tactile sensitivity, a "touch gate", was measured under conditions in which the location of the noxious stimuli was varied with respect to the tactile stimulus applied to the thenar eminence of humans. Vibrotactile thresholds were measured in the absence of pain and during administration of a painful stimulus, with the stimulus frequencies selected to activate independently the four psychophysical channels hypothesized to exist in human glabrous skin. Heat-induced pain produced by spatially co-localizing the noxious stimuli with the tactile stimuli was found, on average, to elevate threshold amplitude by 2.2 times (6.7 dB). Co-localized, cold-induced pain raised the average thresholds by about 1.5 times (3.6 dB). Heat-induced pain presented contralaterally produced no change in vibrotactile sensitivity indicating that the effect is probably not due to attentional mechanisms. Ipsilateral heat-induced pain caused an elevation in tactile thresholds even when the noxious and non-noxious stimuli were not co-localized, and the effect may seem to require that the painful stimulus be within the somatosensory region defined possibly in terms of dermatomal organization. Thus the effect is probably related to somatotopic organization and is not peripherally mediated. A brief discussion as to the possible locus of the touch gate within the nervous system is also given.

Light- and electron-microscopic immunocytochemistry (ICC) was performed on Pacinian corpuscles (PCs) obtained from cat mesentery to determine the presence and location of various proteins within the accessory capsule and the neurite. Antibodies to tubulin, neurofilament 200, actin, collagen II and V, glial fibrillary acidic protein (GFAP) and S-100 were used. Type II collagen was localized only in the outer core of the accessory capsule, which is composed of an inner core, an intermediate layer or growth zone, an outer core and an external capsule. Type V collagen was found only in the intermediate growth zone. Intermediate filaments labeled with anti-GFAP were only found in the inner core. The calcium-binding protein that was labeled by anti-S-100 was found only in the inner core. Diffuse and variable staining for actin is present throughout the accessory capsule. The differences in distribution of these various proteins within the capsule suggest different structural/functional properties of the various capsule regions. The neurite was found to contain microtubules (i.e., tubulin) and neurofilaments throughout, but these cellular inclusions were not found within the cytoplasmic extensions (filopodia) that project from the neurite into the hemilamellar clefts formed by the inner-core hemilamellae. The extensions, however, were found to contain actin in a much greater density than that seen in the neurite proper. The presence of actin, but apparent lack of other cytostructural elements within the extensions, is highly reminiscent of the composition of stereocilia found on vestibular and auditory hair cells. Since stereocilia have been shown to play a role in hair-cell mechanotransduction, it is possible that the cytoplasmic extensions are significantly involved with mechanotransduction within the PC.

Previous experiments performed on monkey and human fingertips suggested that the skin surface and stimulus probe decouple for sinusoidal displacements applied perpendicularly to the skin surface. From these observations, it was concluded that sinusoidal vibration may not be a suitable stimulus for understanding and modeling the tactile system. We repeated these experiments on human observers using stimulus frequencies ranging from 0.5 to 240 Hz and with displacement amplitudes up to 1 mm peak-to-peak (p-p). The skin and probe movements were measured in the steady-state using stroboscopic illumination and video microscopy. Contrary to previous conclusions, we found that decoupling did not occur for amplitudes less then 0.25 mm p-p, regardless of stimulus frequency. Decoupling was only observed for stimulus amplitudes greater than 0.25 mm over the stimulus-frequency range investigated. To further investigate this effect, a modified stimulus contactor was used, which permitted the measurement of the skin's movement using reflected light. Measurements were made on both the index fingertip and the thenar eminence. Regardless of body site, no decoupling between the skin and stimulus probe was observed for frequencies ranging from 20 to 100 Hz up to displacements of 0.25 mm p-p. These levels are well within the range used in most human psychophysical experiments performed on these parts of the body. We conclude that sinusoidal vibration can be used reliably to stimulate the tactile system and is an appropriate stimulus for developing models of touch.

A series of experiments are described in which magnitude estimates of the perceived size of steel balls were made when the balls were actively rolled between the fingertip and several other body sites (thumb, thenar eminence, forearm). This movement, called scripting, involves actively-moving an object by a touching surface over another surface of the body which is passively being touched. We define this active/passive activity as "intra-active touch" and the results show that the perceptual size of the balls is dependent upon the body part passively being activated. An additional series of experiments decoupled the actively generated and passively received tactile information by having subjects either perform the scripting on another individual's body site or by having the other individual roll the balls on the subject's various sites. The latter experiments showed that the passive body can contribute to the overall impression of the size of the balls, but only when the intra-active touching involved the glabrous skin of the hands. Intra-active touch between the active finger and the passively touched hairy skin of the forearm showed no effect of the touched surface on the perceived size of the balls. The results suggest that the mechanisms of intra-active touch are different when glabrous skin activates glabrous skin than when glabrous skin activates hairy skin.

Temporal summation, a decrease in the detection threshold that occurs when either the duration of a stimulus or the number of stimuli in a sequence is increased, has been attributed to the operations of either the mechanism of neural integration or of probability summation. Our experiments indicate that under certain conditions, both mechanisms may operate, but that the process of neural integration is an exclusive characteristic of the Pacinian (P) channel. The P channel was isolated by applying 250 Hz stimuli through a 1.5 cm2 contactor to the thenar eminence of the hand and the NPII channel was isolated by applying the stimuli through a 0.01 cm2 contactor. The finding that the slopes of the psychometric functions were the same within both channels indicated that probability summation could not account for temporal summation for stimulus durations less than 1 s. The finding that the threshold for the detection of multiple-pulse stimuli increased as the interpulse interval increased indicated that, for time intervals less than 800 ms, temporal summation results from neural integration. But for interstimulus intervals greater than 800 ms, probability summation accounts for temporal summation.

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.

A series of experiments demonstrated the role of higher level cognitive processes, such as attention, in tactile perception. The first series of experiments demonstrated that automatic orienting to a tactile stimulus resulted in inhibition of subsequent stimuli at that body site--inhibition of return (IOR). A possible explanation suggests that inhibition of saccades to a body site can cause the inhibition of subsequent stimuli presented to that same site. In contrast, when the subjects strategically oriented attention to the stimulus, the processing of subsequent stimuli at that body site was facilitated. In both of these experiments the skin received exactly the same test stimuli, (100 Hz sine wave presented for 50 ms), but very different effects were observed depending upon attentional strategy. Experimental manipulations showed that this cannot be due to a peripheral masking of the receptors after cue presentation to the target. Rather the results may be explained centrally by cognitive, particularly attentional mechanisms. Cross-modal interactions suggest that tactile processes are facilitated when vision is oriented to the body site receiving stimulation. Possible explanations come from recent findings of spatiotopic maps of different sensory modalities in the superior colliculus of the midbrain and in the parietal lobe. These are integrated with motor systems that control saccades and head orientation towards sensory inputs. Excitatory links among these maps could be the source of the observed facilitation effects.

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.

Several aspects of tactile, thermal and pain perception were evaluated in an individual (R.S.) with a hemorrhagic lesion centered in her left lateral thalamus. Over a 4-year period, psychophysical evaluations were undertaken every 6-8 months, and five magnetic resonance (MR) studies were conducted. Early tests (1991-1992) revealed large contralateral deficits in R.S.'s perception of touch, innocuous temperature, and mechanically evoked cutaneous pain--more so for the upper versus the lower extremity. R.S. showed a similar pattern for heat pain sensitivity, but a more modest deficit than for mechanically evoked pain. She showed a deficit for cold pain sensitivity on her foot, but not for her hand. Thresholds for all types of stimuli ipsilateral to the lesion were within a normative range. Late in 1993, R.S. demonstrated improvements in sensory capacity for touch and mechanically evoked pain contralaterally, although deficits were still evident. During the same period, heat pain sensitivity improved contralaterally, and strikingly, a permanent, ipsilateral hypersensitivity to heat pain developed in her hand. Throughout the entire testing period, R.S.'s ratings of perceived unpleasantness matched the patterns of perceived pain intensity. Thus, the discriminative and the affective dimensions of her pain would change in tandem. However, perceptible innocuous thermal stimuli evoked no affective response when applied contralaterally, despite being described as pleasant when presented ipsilaterally. Throughout the testing period, R.S. reported a persistent numbness on her right hemi-body. Only during a 3-month period in 1995 did she experience spontaneous pain, which was referred to her right foot. The only change in psychophysical performance related to her right foot was a transient but intense thermal allodynia several months prior to her spontaneous pain. The MR studies over this 4-year period showed changes in the extent of edema, gliosis and/or ischemia that could be related to perceptual changes. Thus, the conspicuous observations in this thalamic lesion case were: (i) differential effects upon the various pain modalities (mechanical, heat and cold); (ii) development of thermal allodynia without mechanical allodynia, including an ipsilateral effect; (iii) a deficit in positive affective responses to temperature; and (iv) the different time courses for changes in evoked somesthetic capacity versus spontaneous paresthesias and pathological pain.

Detection thresholds and intensity-difference thresholds were measured on four subjects ranging in age from 19 to 22 years. The stimuli were 250-Hz bursts of vibration applied through a 3.0 cm2 contactor to the thenar eminence of the right hand. Detection thresholds were substantially higher at 20 degrees C than at 30 degrees or 40 degrees C and were only slightly higher at 40 degrees C than 30 degrees C. When the intensity-difference threshold was expressed in relative terms, as the proportion by which two stimuli must differ in amplitude to be discriminated (delta phi/phi), discrimination capacities were unaffected by surface-skin temperature. The results are consistent with the hypothesis that surface-skin temperature alters the sensitivity of tactile receptors, and that, because of the 'near miss' to Weber's law, the relative difference threshold is unaffected substantially by skin temperature. It was concluded that, at least a partial explanation of the 'near miss' lies in the fact that, at low to moderate sensation levels, the P channel is exclusively activated whereas, at moderate to high sensation levels, because of recruitment of activity in Non-Pacinian channels, neural information for intensity discrimination is additionally provided by channels with superior discriminative capacities.

The effectiveness of using vibrotactile threshold measures to aid in the diagnosis of carpal tunnel syndrome (CTS) was evaluated. Thresholds for detecting 1-, 10-, and 300-Hz vibratory stimuli were measured on the fingertips of 24 CTS patients and 20 healthy control subjects. There were no significant differences in threshold for 1- and 300-Hz between the two groups. Although there were significant differences for 10-Hz stimuli, the mean patient threshold was within 1 standard deviation of the mean threshold for the control group. These results indicate that threshold testing is not a suitable diagnostic tool for CTS. Additionally, we examined whether thresholds were elevated in the presence of pain. Seven patients reported experiences of pain and no pain sessions. No significant differences in threshold were found between the two pain conditions, indicating that the presence of pain related to CTS does not affect threshold.

Vibrotactile thresholds were measured on the thenar eminence and the volar forearm at different static depths of skin indentation. Three stimulus frequencies (1, 20, and 200 Hz) were delivered through either a 0.008- or a 2.9-cm2 contractor. The indentation depths ranged from 0 to 1 mm (0.25-mm steps) relative to the point of skin contact with the stimulator. There was a significant effect of indentation in all stimulus combinations of contactor size, location, and frequency. These results resolve an apparent discrepancy in the literature regarding threshold reduction with increasing contactor size observed on the forearm at low frequencies.

Vibrotactile forward masking experiments were conducted on the thenar eminence of the hand to determine the time course of masking in the Pacinian (P) or a non-Pacinian (NP I) channel. Brief masking and test stimuli that contained energy centered either below 27 or at 500 Hz were used to preferentially activate the NP I or P channels, respectively. Test thresholds were measured at different fixed delays ( Delta t) between the masking and test stimuli that ranged from 5 to 995 microseconds. Masking stimulus level was also varied from 5 to 25 dB above threshold. The masking effect followed an exponential decay with different time constants for each of the channels, but a similar asymptote (residual masking) for both channels. At the higher masking stimulus levels, the time constant for the P channel was about 40 microseconds, while the time constant for the NP I channel was about 100 microseconds. Residual masking in both channels increased about 1 dB for every 5-dB increase of masking stimulus level.

Electron-microscopic analyses of the mitochondrial organization within the neurite innervating the Pacinian corpuscle (PC) were performed to test the hypothesis that the sites of mechanotransduction are the filopodia projecting from the neurite's surface. Since high concentrations of mitochondria imply the need for metabolic energy, and since transduction mechanisms are heavily dependent on such energy, it was reasoned that the greatest concentration of mitochondria should occur near the filopodia if they are involved in mechanotransduction. The analysis that the mitochondria lie close to the terminal neurite's membrane, on average within 0.4 microns, and thus are ideally located for supplying energy for membrane mechanisms. Although they can be found in a ring-like array, as seen in cross-sections of the terminal neurite, their greatest concentration occurs where the filopodia project from the terminal neurite. A linear algebraic analysis of the data set confirmed a high probability of the joint occurrence of a filopodial base and an increased frequency (number) of mitochondria. Thus the results provide further circumstantial support for the hypothesis that transduction within the PC mechanoreceptors takes place at or near the filopodia.

Detection thresholds and difference limens were measured for 16 subjects ranging from 19 to 91 years of age. The stimuli were 250-Hz bursts of vibration applied through a 3.0-cm2 contactor to the thenar eminence of the right hand. Detection thresholds were higher in older than in younger subjects, as were the absolute values of difference limens. When the difference limen was expressed in relative terms as the proportion by which two stimuli had to differ in amplitude to be discriminated (delta alpha/alpha), discriminative capacities were unaffected by aging except for stimuli slightly above the detection threshold, in which case the limens of older subjects were significantly higher than those of younger subjects. The results are consistent with the hypothesis that elevations in the detection thresholds of older subjects are the results of reduced afferent input to central brain centers that, with regard to their capacity to detect the presence of threshold-level stimuli and to discriminate differences among suprathreshold stimuli, are relatively unaffected by aging.

Tactile thresholds for detecting a 50-ms signal presented 25 ms after the termination of a masking stimulus increased as a function of the amplitude level and duration of the masking stimulus. The effects were similar in both the P and NP I channels measured at 250 and 20 Hz, respectively. It was concluded that the increased masking caused by increasing the duration of the masking stimulus resulted from processes other than or in addition to temporal integration--the latter being a characteristic of the P, but not the NP I, channel. The slopes of the masking functions, in which threshold shifts were plotted as a function of masking-stimulus sensation level, were consistently greater for 20-Hz than for the 250-Hz stimuli, suggesting that masking efficiency is greater in the NP I than in the P channel.

Previous reports have indicated that certain mouthrinses, even when used as directed can induce oral pain. In order to help determine the causal agent(s), various commercially available mouthrinses, as well as control solutions, were tested in a psychophysical study in which subjects rated categories of pain during and after mouthrinsing. More specifically, the studies tested the effects of ethanol concentration on induced pain. The results show that there is a direct relationship between ethanol content and the amount of induced pain. Furthermore, the amount of pain was found to increase with time of rinsing, and to slowly decrease after cessation of rinsing. Lastly, comparison of ethanol/water controls with a marketed product (Clear Choice) matched for ethanol content showed that, while ethanol was the key factor in mouthwash-induced oral pain, other presently unidentified agents can also add to the effect.

In this study, the hypothesis that the difference limen (DL) for the detection of differences in amplitude of vibrotactile stimuli is independent of the slope of the sensation magnitude function was tested. The slope of the sensation magnitude function was varied by presenting test stimuli in the presence of or in the absence of vibrotactile noise. The slopes of the sensation magnitude functions were determined through a matching technique in which the subject adjusted stimulus amplitudes of a 250-Hz stimulus presented alone and a 250-Hz stimulus presented simultaneously with a masking noise, so that their sensation magnitudes were equated. The slope of the matching function was found to increase as a function of the intensity of the masking noise. In the second phase of the experiment, the amplitude DL was measured by the gated-pedestal method for test stimuli presented under the same stimulus conditions as used in the matching procedure. At all levels of stimulus intensity, the DL was found to be independent of the masking condition provided the sensation magnitudes of the stimuli were the same. This finding supports the hypothesis that the size of the DL is independent of the slope of the sensation magnitude function, provided the sensation magnitudes of stimuli are the same. The generality of this principle, first discovered in hearing, is thus extended to another sense modality.

A hybrid digital/analog device capable of making high-resolution linear and area measurements from a standard monochromatic video image is described. The device is capable of dynamic as well as static data acquisition when used in conjunction with standard NTSC video recording equipment. Digital output allows for computer interfacing. Linear dimensions are obtained by electronically superimposing two horizontal and two vertical scaling lines on a video monitor. Each linear dimension has an eight-bit resolution and is displayed on the front panel with seven segment LED's. Eight-bit, digital-to-analog converters are also used to provide analog outputs. A measurement of the temporal deformation pattern of the accessory capsule surrounding a mechanoreceptor (Pacinian corpuscles) in response to vibratory displacements is demonstrated. Area measurements are obtained via a window comparator, a 6-MHz clock, and a 16-bit digital-to-analog converter. Data is only valid within the zone set by the scaling lines, allowing data regions to be isolated from noise, etc. The 16-bit digital signal measuring the area of interest is then converted for analog output. The measurement of the consensual pupillary reflex in response to full-field illumination (Ganzfeld) is given as an example.

Thresholds for the detection of vibrotactile signals of varied duration applied to the thenar eminence were measured in the absence of and in the presence of a masking stimulus. Signals were 250- and 500-Hz sinusoids and noise bursts with bandwidth limited to 250-1000 Hz. The masking stimulus was either a 250-Hz sinusoid, which was presented in phase with the signal when it was sinusoidal, or noise. Changes in threshold as a function of changes in signal duration were found which were predicted accurately from Zwislocki's theory [J. Acoust. Soc. Am. 32, 1046-1060 (1960)] of temporal summation when the signal was detected by the Pacinian channel, but not when it was detected by a non-Pacinian channel (NP). However, when either the signal or the masking stimulus or both were noise, NP thresholds were affected by changes in signal duration. Only when the signal and masking stimuli were both sinusoids were NP thresholds independent of signal duration. It is concluded that signal duration effects in the NP channel are not due to temporal integration, but rather to increases in information about the signal content provided to the subject as exposure duration is increased.

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.