Neural Mechanisms of Absolute Tactile Localization in Monkeys

Macaca nemestrina monkeys were trained to indicate the location of suprathreshold tactile stimuli delivered to the glabrous skin of either foot. The testing paradigm involved self-initiated trials (a bar press), followed by 10-Hz stimulation at one of six locations (e.g., on the distal phalanx of the second toe on the left foot), providing the opportunity for the animal to press one of six buttons located on a facing panel. The buttons were positioned on a picture of a monkey's feet at locations corresponding to the skin loci that were stimulated on different trials. If the animal first presed the button corresponding to the position stimulated, liquid reward was delivered; responses to any other button terminated stimulation without reward, requiring initiation of another trial for the opportunity to receive reinforcement.

The localization errors for normal monkeys were reliably greater along the mediolateral dimension of the foot than they were proximodistally. For example, stimulation of the tip of toe 4 elicited responses to the button at the tip of toe 2 on 25% of the trials, as compared with only 10% errors betweeen the tip of toe 4 and the pad at the base of toe 4. Following unilateral interruption of the dorsal spinal columns at an upper thoracic level, the capacity for absolute tactile localization was unchanged over months of testing. The greater localization accuracy along the proximodistal axis of the foot remained after dorsal column transection.

In order to evaluate neural substrates of localization by monkeys, single-neuron receptive field (RF) sizes and distributions within the first somatosensory (SI) cortex were examined to determine the overlap or separation of the representations of different points on glabrous skin. The sample of neurons that provided the RF data was obtained in previous investigations of unanesthetized, neuromuscularly blocked Macaca fascicularis monkeys. Analysis of RF overlap revealed that greater than 50% of cytoarchitectural area 1 units that responded to stimulation of one digit tip also responded to another digit or to the pad at the base of a digit. These large RFs seem poorly suited to subserve a high degree of spatial localization and are compatible with the frequent localization errors by the monkeys in the behavioral experiments. However, the area 1 RF data do not explain the tendency of these animals to exhibit better localization accuracy along the proximodistal axis than along the mediolateral axis of the volar foot. We suggest that the observed asymmetry of localization accuracy may be the result of dynamic neural mechanisms that involve lateral interactions between cortical columns. Evidence from metabolic mapping experiments suggests that these lateral interactions determine the spatial distribution of cortical columns that respond to repetitive somatic stimuli (Whitsel et al, 1988).     

Somatotopy of digital nerve projections to the dorsal horn in the monkey

The dorsal horn projection patterns of finger nerves were investigated in four Macaca mulatta monkeys. Proper digital branches of the median nerves, serving the radial aspect of a digit on each hand, were loaded with wheatgerm agglutinin-horseradish peroxidase complex (WGA:HRP). The distribution of the lectin-enzyme complex was mapped in the right and left dorsal horns. The dorsal horn projections of the digital nerves were localized in segments C6-C8 in laminae I-VI, primarily in laminae I-IV. The wedge-shaped termination zones were somatotopically organized, in agreement with the projections of the digits in cats. The fingers are represented medially, as they are in the cat. This similarity suggests that there is a mediolateral gradient of dorsal horn organization similar to that of the cat, with distal skin represented medially and proximal skin represented laterally. The rostrocaudal trajectory of finger representation, with digit 1 most rostral and digit 5 most caudal, is also in agreement with the organization of hindlimb toe projections in the cat. There was a high degree of bilateral symmetry for homologous nerves, and little overlap of projections from nerves innervating adjacent fingers. The sample size was too small to permit us to assess interanimal variation. These results suggest a similar somatotopy of projections, and presumably of dorsal horn cell somatotopy, in monkey and cat.     

Evidence for polarizing zone in the limb buds of Xenopus laevis

To test for the presence of polarizing mesoderm in an amphibian, Xenopus laevis hindlimb bud tips were rotated 180° on the proximodistal axis and returned to the stump. Supernumerary outgrowths were induced in the preaxial stump and preaxial tip tissues, and the most postaxial digit always formed next to the grafted postaxial tissue. The occurrence of polarized supernumerary outgrowths indicated that the posterior limb border contained a polarizing zone. When the limb tip was cut at varying known lengths from the body wall, rotated, and grafted to the limb stump, the incidence of twinning along the proximodistal axis permitted insight into the distribution of the polarizing zone along the posterior border. The location of polarizing tissues was found to be similar to that in the chick wing bud at comparable stages. To confirm the posterior border stump influence on the rotated preaxial limb tip tissues, 180° tip rotations were made at the proximodistal level with the highest incidence of twinning. In these cases, the adjacent stump posterior border tissues (polarizing zone) were removed, leaving a substantial amount of the deeper postaxial stump tissue, however. The frequency of twinning from tip tissues was greatly reduced in these larvae compared to those with rotated limb tips on intact stumps. Cytological examination of supernumerary outgrowths resulting from grafts of two-nucleolate tips onto one-nucleolate stumps confirmed the preaxial source of the supernumerary outgrowths.     

Somatotopy of Digital Nerve Projections to the Dorsal Horn in the Monkey

The dorsal horn projection patterns of finger nerves were investigated in four Macaca mulatto monkeys. Proper digital branches of the median nerves, serving the radial aspect of a digit on each hand, were loaded with wheatgerm agglutinin—horseradish peroxidase complex (WGA:HRP). The distribution of the lectin—enzyme complex was mapped in the right and left dorsal horns.

The dorsal horn projections of the digital nerves were localized in segments C₆-C₈ in laminae I-VI, primarily in laminae I-IV. The wedge-shaped termination zones were somatotopically organized, in agreement with the projections of the digits in cats. The fingers are represented medially, as they are in the cat. This similarity suggests that there is a mediolateral gradient of dorsal horn organization similar to that of the cat, with distal skin represented medially and proximal skin represented laterally. The rostrocaudal trajectory of finger representation, with digit 1 most rostral and digit 5 most caudal, is also in agreement with the organization of hindlimb toe projections in the cat. There was a high degree of bilateral symmetry for homologous nerves, and little overlap of projections from nerves innervating adjacent fingers. The sample size was too small to permit us to assess interanimal variation. These results suggest a similar somatotopy of projections, and presumably of dorsal horn cell somatotopy, in monkey and cat.     

Effect of Mechanical Stimulus Spread across Glabrous Skin of Raccoon and Squirrel Monkey Hand on Tactile Primary Afferent Fiber Discharge

The role of spread of skin deformation in activating cutaneous mechanoreceptors at a distance from their threshold receptive fields (RFs) was examined in glabrous skin of the North American raccoon and the squirrel monkey. One feedback-controlled mechanical stimulus probe was used to indent the skin to a controlled depth at a constant velocity, at varying distances from a second probe, which was used to monitor vertical displacement depth and velocity at this distant site. In many instances, the monitor probe was positioned over the RF of a cutaneous mechanoreceptor, and single-unit action potentials were simultaneously recorded from individual fibers of the median or ulnar nerve.

With distance from the site of stimulation, there was a systematic, monotonic decline in indentation depth and velocity; velocity fell off with distance more rapidly than depth. The degree of diminution with distance varied with the size, shape, and curvature of the digital or palm pad stimulated. Spread of indentation was more restricted on digital than on palm pads, and was more restricted across monkey skin than across raccoon skin. Spread was less with higher-velocity than with lower-velocity indentations, but was seemingly unaffected by indentation depth.

As expected from the findings noted above, the number of spikes discharged by slowly adapting mechanoreceptive afferent fibers declined more rapidly with distance between stimulus site and RF for digital than for palmar RFs, in squirrel monkey than in raccoon skin, and with higher-velocity than with lower-velocity stimuli. Furthermore, the number of spikes occurring during either ramp or early static indentation phases of stimulation dropped to zero more rapidly with distance than did either vertical indentation depth or velocity. Decreases with distance in both indentation depth and velocity acted to restrict the size of suprathreshold RFs. For most units, horizontal components of mechanical stimulation subtracted from the effects of vertical components.

It is suggested, on the basis of this and other studies, that many neural and perceptual phenomena usually attributed to central mechanisms of afferent inhibition may be attributable, at least in part, to mechanical properties of the skin. In addition, the present data suggest that regional variations in the two-point limen may be associated with variations in spread of mechanical deformation. The conclusion that glabrous skin and subjacent soft tissues act as a low-pass filter system provides a mechanical basis for the relative efficacy of high-frequency vibratory stimuli in tactile pattern perception. Finally, the view is presented that the skin and subjacent tissues should be considered, along with cutaneous mechanoreceptors, as forming a tactile receptor organ system.     

Localization of Merkel cells in the monkey skin: an anatomical model

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.     

Experience-induced plasticity of cutaneous maps in the primary somatosensory cortex of adult monkeys and rats

In a first study, the representations of skin surfaces of the hand in the primary somatosensory cortex, area 3b, were reconstructed in owl monkeys and squirrel monkeys trained to pick up food pellets from small, shallow wells, a task which required skilled use of the digits. Training sessions included limited manual exercise over a total period of a few hours of practice. From an early clumsy performance in which many retrieval attempts were required for each successful pellet retrieval, the monkeys exhibited a gradual improvement. Typically, the animals used various combinations of digits before developing a successful retrieval strategy. As the behavior came to be stereotyped, monkeys consistently engaged surfaces of the distal phalanges of one or two digits in the palpation and capture of food pellets from the smallest wells. Microelectrode mapping of the hand surfaces revealed that the glabrous skin of the fingertips predominantly involved in the dexterity task was represented over topographically expanded cortical sectors. Furthermore, cutaneous receptive fields which covered the most frequently stimulated digital tip surfaces were less than half as large as were those representing the corresponding surfaces of control digits. In a second series of experiments, Long-Evans rats were assigned to environments promoting differential tactile experience (standard, enriched, and impoverished) for 80 to 115 days from the time of weaning. A fourth group of young adult rat experienced a severe restriction of forepaw exploratory movement for either 7 or 15 days. Cortical maps derived in the primary somatosensory cortex showed that environmental enrichment induced a substantial enlargement of the cutaneous forepaw representation, and improved its spatial resolution (smaller glabrous receptive fields). In contrast, tactile impoverishment resulted in a degradation of the forepaw representation that was characterized by larger cutaneous receptive fields and the emergence of non-cutaneous responses. Cortical maps derived in the hemispheres contralateral to the immobilized forelimb exhibited a severe decrease of about 50% in the overall areal extent of the cutaneous representation of the forepaw, which resulted from the invasion of topographically organized cortical zones of non-cutaneous responses, and numerous discontinuities in the representation of contiguous skin territories. The size and the spatial arrangement of the cutaneous receptive fields were not significantly modified by the immobilization of the contralateral forelimb. Similar results were obtained regardless of whether the forelimb restriction lasted 7 or 15 days. These two studies corroborate the view that representational constructs are permanently reshaped by novel experiences through dynamic competitive processes. These studies also support the notion that subject-environment interactions play a crucial role in the maintenance of basic organizational features of somatosensory representations.     

Tactile rivalry demonstrated with an ambiguous apparent-motion quartet

When observers view ambiguous visual stimuli, their perception will often alternate between the possible interpretations, a phenomenon termed perceptual rivalry [1]. To induce perceptual rivalry in the tactile domain, we developed a new tactile illusion, based on the visual apparent-motion quartet [2]. Pairs of 200 ms vibrotactile stimuli were applied to the finger pad at intervals separated by 300 ms. The location of each successive stimulus pair alternated between the opposing diagonal corners of the approximately 1 cm(2) stimulation array. This stimulation sequence led all participants to report switches between the perception of motion traveling either up and down or left and right across their fingertip. Adaptation to tactile stimulation biased toward one direction caused subsequent ambiguous stimulation to be experienced in the opposing direction. In contrast, when consecutive trials of ambiguous stimulation were presented, motion was generally perceived in the direction consistent with the motion reported in the previous trial. Voluntary eye movements induced shifts in the tactile perception toward a motion axis aligned along a world-centered coordinate frame. Because the tactile quartet results in switching perceptual states despite unvaried sensory input, it is ideally suited to future studies of the neural processes associated with conscious tactile perception.     

Acute Effects of Total or Partial Digit Denervation on Raccoon Somatosensory Cortex

The immediate effects of total or partial denervation of single digits (0-16 hr after nerve transection) on primary somatosensory cortex were studied electrophysiologically. Comparisons of response properties and cortical somatotopy were made between intact raccoons and four groups of raccoons with transection of some or all of the nerves innervating the fourth or fifth digit. Animals with all four digital nerves cut (amputation of the digit) were most different from normal. Approximately half of the penetrations in the affected cortical region showed inhibitory responses to stimulation of adjacent skin regions. These consisted of a strong response to stimulus offset and/or a suppression of spontaneous activity during indentation. Since these responses were substantially different from those recorded several months after digit amputation, additional changes in connectivity and synaptic strength must occur with chronic denervation. These inhibitory responses were not seen in animals with one, two, or three nerves cut per digit.

In the animals with partial denervation of a digit, the greatest disruption occurred when both ventral nerves to the glabrous skin were transected. This yielded cell clusters with abnormally large receptive fields, disruptions in somatotopic organization, and a decreased occurrence of low-threshold responses. If only one nerve to glabrous skin was transected, there was less change, even if it was combined with transection of both nerves to hairy skin. These results suggest that the release of inhibitory responses in a cortical digital region by amputation is prevented by the retention of even one ventral nerve. None of the denervation conditions produced large nonresponsive areas of cortex, which would have indicated a loss of all inputs.     

Acute effects of total or partial digit denervation on raccoon somatosensory cortex

The immediate effects of total or partial denervation of single digits (0-16 hr after nerve transection) on primary somatosensory cortex were studied electrophysiologically. Comparisons of response properties and cortical somatotopy were made between intact raccoons and four groups of raccoons with transection of some or all of the nerves innervating the fourth or fifth digit. Animals with all four digital nerves cut (amputation of the digit) were most different from normal. Approximately half of the penetrations in the affected cortical region showed inhibitory responses to stimulation of adjacent skin regions. These consisted of a strong response to stimulus offset and/or a suppression of spontaneous activity during indentation. Since these responses were substantially different from those recorded several months after digit amputation, additional changes in connectivity and synaptic strength must occur with chronic denervation. These inhibitory responses were not seen in animals with one, two, or three nerves cut per digit. In the animals with partial denervation of a digit, the greatest disruption occurred when both ventral nerves to the glabrous skin were transected. This yielded cell clusters with abnormally large receptive fields, disruptions in somatotopic organization, and a decreased occurrence of low-threshold responses. If only one nerve to glabrous skin was transected, there was less change, even if it was combined with transection of both nerves to hairy skin. These results suggest that the release of inhibitory responses in a cortical digital region by amputation is prevented by the retention of even one ventral nerve. None of the denervation conditions produced large nonresponsive areas of cortex, which would have indicated a loss of all inputs.     

Branching,segmentation and the metapterygial axis: pattern versus process in the vertebrate limb

Explanations of the patterns of vertebrate fin and limb evolution are improving as specific hypotheses based on molecular and developmental data are proposed and tested. Comparative analyses of gene expression patterns and functions in developing limbs, and morphological patterns in embryonic, adult and fossil limbs point to digit specification as a key developmental innovation associated with the origin of tetrapods. Digit development during the fin-to-limb transition involved sustained proximodistal outgrowth and a new phase of Hox gene expression in the distal fin bud. These patterning changes in the distal limb have been explained by the linked concepts of the metapterygial axis and the digital arch. These have been proposed to account for the generation of limb pattern by sequential branching and segmentation of precartilagenous elements along the proximodistal axis of the limb. While these ideas have been very fruitful, they have become increasingly difficult to reconcile with experimental and comparative studies of fin and limb development. Here we argue that limb development does not involve a branching mechanism, and reassess the concept of a metapterygial axis in limb development and evolution.     

Tactile discrimination of gratings: psychophysical and neural correlates in human and monkey.

Human and monkey performance on discriminating tactile gratings revealed comparable cross-species Weber functions. Neural data obtained while monkeys performed discriminations revealed some matching of neural and psychometric functions. Nearly constant firing rate differences occurred at discrimination threshold for unequal groove widths. Firing rate differences of some cells decreased on trials preceding discrimination errors, and thus predicted performance.     

Tactile Discrimination of Gratings: Psychophysical and Neural Correlates in Human and Monkey

Human and monkey performance on discriminating tactile gratings revealed comparable cross-species Weber functions. Neural data obtained while monkeys performed discriminations revealed some matching of neural and psychometric functions. Nearly constant firing rate differences occurred at discrimination threshold for unequal groove widths. Firing rate differences of some cells decreased on trials preceding discrimination errors, and thus predicted performance.     

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.     

Relations between Stimulus Force,Skin Displacement,and Discharge Characteristics of Slowly Adapting Type I Cutaneous Mechanoreceptors in Glabrous Skin of Squirrel Monkey Hand

(1) The contributions of viscoelastic properties of squirrel monkey glabrous skin to slowly adapting Type I (SAI) mechanore-ceptive afferent fiber discharge were examined in the present study. Individual fibers of the median and ulnar nerves were isolated by microdissection in six monkeys anesthetized with pentobarbital sodium. Utilizing mechanical stimulation and data analysis techniques identical to those of a previous study of raccoon glabrous skin and its mechanoreceptors (Pubols, 1982a; Pubols and Maliniak, 1984), we studied and compared responses to punctate mechanical stimuli controlled with respect to force or displacement. (2) Squirrel monkey glabrous skin was found to be more compliant than raccoon glabrous skin, in that a given force applied to either a digital or a palmar skin pad produced a greater displacement of squirrel monkey skin. Skin displacement increased approximately linearly with increasing forces at the beginning of static stimulation, but over time (at least up to 20 sec), the relationship became negatively accelerated. (3) Absolute-force thresholds of individual SAI units were significantly lower in squirrel monkey (mean = 122 mg, range = 48-340 mg) than in raccoon (mean =484 mg, range = 70-1,290 mg). However, absolute-displacement thresholds were insignificantly lower (squirrel monkey: mean = 17.24 μm, range = 5-30μ raccoon: mean = 30 μm, range = 5-185 μm). (4) Application of suprathreshold forces (range = 1-20 g) and displacements (range = 500-1,000 μm) revealed greater interunit variability in response to maintained stimulation than previously found in raccoon. In 8 out of 15 fibers, the rate of adaptation was significantly greater during constant-displacement than during constant-force stimulation; in 4 cases there was no significant difference; and in 3 cases the rate of adaptation was significantly greater during constant-force stimulation. (5) Potential sources of interunit variability include surface topography of the hand, properties of cutaneous and subcutaneous tissues in the vicinity of the receptor, and experimental variables such as stimulus amplitude and rate of stimulus onset. (6) It is suggested that both regional and species differences in functional properties of cutaneous mechanore-ceptors are more likely attributable to differences in mechanical properties of skin and subjacent tissues than to any inherent differences in receptor properties.     

Spatial features of vibrotactile masking effects on airpuff-elicited sensations in the human hand

It was recently shown that the cutaneous sensitivity to airpuffs is decreased by a low-frequency vibrotactile masker in the hairy skin, and by a low-frequency but especially by a high-frequency masker in the glabrous skin. In the current study, the spatial features of this masking effect were determined in four healthy human subjects, using a reaction time paradigm. The masking effect decreased monotonically with increasing interstimulus distance, and identically in longitudinal and transverse (i.e., lateral) directions in the palm or dorsal surface of the hand. The masking effect was stronger in the glabrous than in the hairy skin, especially in the fingers. In the glabrous skin, the spread of masking effect produced by a high-frequency masker was more extensive than that produced by a low-frequency masker. The mechanical spread of high-frequency vibration was less extensive than that of low-frequency vibration in the skin. In the glabrous skin, a masker applied to the tip of the finger produced a stronger masking effect on sensations in the base of the finger than when the masker was located at the base and the test stimulus was located at the tip. It is concluded that mechanical spread of vibration in the skin is of minor importance in explaining the masking effects. Different peripheral neural mechanisms underlie the airpuff-elicited sensations in the hairy and glabrous skin. The afferent inhibitory mechanisms are stronger for signals coming from the glabrous skin of the fingers than for signals coming from the hairy skin. Furthermore, the peripheral innervation density and size of the cortical representational areas may be of importance in determining the magnitude of the masking effect.     

An experimental unit for the recording of the escape reaction of a ground snail to tactile stimulation

An original working experimental unit for noninvasive objective recording of the magnitude of escape reaction of a ground snail evoked by tactile stimulation is described. A. snail creeps upwards over the cylinder rotating around its horizontal axis. A watching device ensures a constant snail position relative to a light source and a photoelement. A device for tactile stimulation which provides graduated energy of an impact is constructed on the basis of the magnetic circuit of a loudspeaker. In response to a tactile stimulus a snail pulls in its feelers, head, and foot, and the area of snail's shadow decreases. These changes are indicated by the photoelement.     

Spatial Features of Vibrotactile Masking Effects on Airpuff-Elicited Sensations in the Human Hand

It was recently shown that the cutaneous sensitivity to airpuffs is decreased by a low-frequency vibrotactile masker in the hairy skin, and by a low-frequency but especially by a high-frequency masker in the glabrous skin. In the current study, the spatial features of this masking effect were determined in four healthy human subjects, using a reaction time paradigm. The masking effect decreased monotonically with increasing interstimulus distance, and identically in longitudinal and transverse (i.e., lateral) directions in the palm or dorsal surface of the hand. The masking effect was stronger in the glabrous than in the hairy skin, especially in the fingers. In the glabrous skin, the spread of masking effect produced by a high-frequency masker was more extensive than that produced by a low-frequency masker. The mechanical spread of high-frequency vibration was less extensive than that of low-frequency vibration in the skin. In the glabrous skin, a masker applied to the tip of the finger produced a stronger masking effect on sensations in the base of the finger than when the masker was located at the base and the test stimulus was located at the tip. It is concluded that mechanical spread of vibration in the skin is of minor importance in explaining the masking effects. Different peripheral neural mechanisms underlie the airpuff-elicited sensations in the hairy and glabrous skin. The afferent inhibitory mechanisms are stronger for signals coming from the glabrous skin of the fingers than for signals coming from the hairy skin. Furthermore, the peripheral innervation density and size of the cortical representational areas may be of importance in determining the magnitude of the masking effect.     

躯体感觉Ⅱ区皮层内微刺激对猕猴回避性操作式条件反射和针刺镇痛的影响

在七只清醒、可以活动的猕猴上观察了皮层内刺激 S_Ⅱ区对外周痛阈和针刺镇痛的影响,其结果如下:(1)在73次皮层内刺激的实验中有72次引起对侧相应皮肤感受野的痛阈变化,其中54次痛阈明显升高。痛阈升高的效应在停止刺激后常持续0.5—3min。较浅层的刺激,痛阈升高比较明显;不同刺激强度引起的痛阈升高的程度也不相同。(2)一般皮层内刺激 S_Ⅱ区也可导致同侧相应皮肤感受野痛阈升高,但不如对侧痛阈升高明显。(3)皮层内刺激 S_Ⅱ区时,非感受野痛阈几乎没有任何改变。(4)S_Ⅱ区皮层内刺激可增强针刺镇痛效应。