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Conduction velocity of the human spinothalamic tract as assessed by laser evoked potentials.

To study the conduction velocity of the spinothalamic tract (STT) we delivered CO2 laser pulses, evoking pinprick sensations, to the skin overlying the vertebral spinous processes at different spinal levels from C5 to T10 and recorded evoked potentials (LEPs) in 15 healthy human subjects. These stimuli yielded large-amplitude vertex potentials consisting of a negative wave at a peak latency of about 200 ms followed by a positive wave at a peak latency of about 300 ms. The mean conduction velocity of the STT was 21 m/s, i.e. higher than the reported velocity of the corresponding primary sensory neurons (type II AMH). Because dorsal stimulation readily yields reproducible brain LEPs, we expect this technique to be useful as a diagnostic tool for assessing the level of spinal cord lesions.

Adult↗

Ultrastructural analysis of axosomatic contacts on functionally identified primate spinothalamic tract neurons.

The morphology and frequency of axosomatic contacts on three functionally identified primate spinothalamic tract (STT) cells were analyzed at the electron microscopic level. The STT cells analyzed were wide-dynamic-range neurons responsive to activation of low- and high-threshold cutaneous afferents innervating the foot. The somas were located in the lateral border of lamina V; the dendritic trees were oriented dorsally and were very extensive. Numerous spinelike appendages were observed emanating from two of the cell bodies. Terminal types contacting the cell bodies were categorized at several different layers through each neuron. Six morphologically different terminal types were established following analysis of serial sections. Profiles classified as round (R) terminals containing round clear vesicles and zero or one dense-core vesicle made up over 50% of the total population in contact with the STT somas. Profiles containing round clear vesicles and two to four small-diameter dense-core vesicles (D1 category) made up approximately 10% of the population in contact with each soma. Flat (F) terminals with oblong or flattened clear vesicles made up approximately 8% of the population. The remaining three categories (D2, L1, and L2) distinguished by the number and size of the dense-core vesicles made up a small percentage of the total population in contact with the cell bodies. The distribution of terminal types on the soma proper versus somatic spines was also determined for one cell. The proportions of the six terminal types contacting the soma of these cells were very similar, although the physiological characteristics of each cell were different. However, the relative proportions of terminal types on these three lamina V STT cell bodies were different from those previously reported contacting somata in lamina V, suggesting that there may be a unique innervation of STT cells that differentiates them from other cell types in lamina V.

Action Potentials↗

Responses of primate spinothalamic tract neurons to renal pelvic distension.

1. Experiments were performed to examine responses of spinothalamic tract (STT) neurons to distension of the renal pelvis. Nineteen monkeys (Macaca fascicularis) were anesthetized with alpha-chloralose, paralyzed, and artificially ventilated. Fifty-four STT neurons in the T11-L2 segments were studied. Each cell was excited by renal nerve stimulation and had a somatic receptive field in the left flank and/or the abdomen. 2. Distension of the left renal pelvis to 50 mmHg for 20-30 s increased activity of 40 STT neurons. Two types of responses were observed. Six cells responded rapidly to the increase in renal pelvic pressure. Thereafter activity of these cells completely adapted. The other 34 cells also responded rapidly to the distension: however, the subsequent adaptation was not complete. Average activity before distension was 13 +/- 1 (SE) spikes/s. Distension increased activity to a peak of 42 +/- 3 spikes/s. Mean activity just before the end of the distension was 27 +/- 3 spikes/s. 3. The pelvic pressure-cell response relation was determined for 16 cells. Only one cell responded to a pressure of 20 mmHg. Three responded to 30 mmHg, and all others responded to 40 mmHg and higher. The average response threshold was 32 +/- 1 mmHg. Peak responses increased as distending pressure increased from 40-80 mmHg. Responses to a pressure of 100 mmHg were no greater than to 80 mmHg. Adapted levels of activity were also a function of distending pressure in the 40-80 mmHg range. 4. Probability of responses was unrelated to somatic input. However, cells with A delta- and C-fiber renal input were significantly more likely to respond to renal pelvic distension than cells with only A delta-renal input. Magnitude of responses to a pressure of 50 mmHg was not related to the type of renal input to the cells; however, among the cells tested at all pressures, cells with A delta- and C-fiber input had significantly greater responses to pressures of 80 and 100 mmHg. 5. Cells were studied in laminae I and IV-VII: responses were unrelated to laminar location. None of the 6 cells located in L2 responded to renal pelvic distension; 8 of 12 in L1 responded; 24 of 28 in T12 responded; and all 8 cells in T11 responded. 6. Stimulation of inhibitory receptive fields on the right hindlimb reduced activity of four cells to a significantly greater extent during pelvic distension than before pelvic distension.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Distribution of trigeminothalamic and spinothalamic-tract neurons showing substance P receptor-like immunoreactivity in the rat.

Trigeminothalamic and spinothalamic-tract neurons provided with substance P receptor (SPR) were examined in the rat by SPR immunofluorescence histochemistry combined with Fluoro-Gold (FG) fluorescent retrograde labeling. After FG injection in the thalamic regions, FG-labeled cells with SPR-like immunoreactivity were seen mainly in laminae I and III of the medullary and spinal dorsal horns and lateral spinal nucleus. In these regions, about one-fourth to one-third of FG-labeled cells showed SPR-like immunoreactivity.

Afferent Pathways↗

The effect of a kainate GluR5 receptor antagonist on responses of spinothalamic tract neurons in a model of peripheral neuropathy in primates.

The responses of antidromically identified spinothalamic tract (STT) neurons to mechanical and thermal stimuli were compared in anesthetized normal and neuropathic monkeys before and after administration of a GluR5 kainate receptor antagonist (LY382884) into the spinal cord dorsal horn through a microdialysis fiber. Peripheral neuropathy was induced by tight ligation of the L7 spinal nerve 13-15 days prior to the experiment. STT neurons recorded in the animals with neuropathy showed increased responsiveness to weak mechanical stimuli and to heating and cooling of the skin compared to STT cells in normal animals. In both normal and the neuropathic monkeys the responses of the STT neurons to mechanical and thermal stimuli were attenuated by LY382884 application in a concentration-dependent manner. Intraspinal application of LY382884 in the neuropathic animals led to a potent reduction of those responses of the STT neurons that were aggravated by the peripheral neuropathy (weak mechanical, heat and innocuous cooling stimuli). These results suggest that kainate receptors are involved in synaptic activation of STT cells in the normal state and may also play an important role in pathological pain states such as peripheral neuropathy in primates. Kainate receptor antagonists could thus be useful for the treatment of certain forms of allodynia and hyperalgesia.

Animals↗

The cells of origin of the spinothalamic tract of the rat: a quantitative reexamination.

We quantitatively reinvestigated the cells of origin of the spinothalamic tract (STT) of the rat. Injections of Fluoro-Gold that filled the thalamus on one side labeled large numbers of neurons throughout the length of the spinal cord. In 3 cases, we estimated the total number of STT neurons by counting labeled neurons in 18 of the 34 total segments, applying correction factors to these counts, and estimating the numbers of labeled neurons in the 16 remaining unexamined segments. The accuracy of these estimates was tested in two animals in which labeled neurons were counted in all 34 spinal segments. In both cases, the estimated totals of STT neurons differed from the counted totals by less than 5%. In the most effective case, we estimated that more than 9500 STT neurons were labeled. This study indicates that the number of STT neurons in rats is larger than previously reported and suggests that the STT may play an important role in nociception in rats, as it does in primates including humans.

Animals↗

Infusion of substance P or neurokinin A by microdialysis alters responses of primate spinothalamic tract neurons to cutaneous stimuli and to iontophoretically released excitatory amino acids.

The responses of 25 spinothalamic tract (STT) neurons to mechanical and thermal stimulation of the skin, as well as to a battery of iontophoretically applied excitatory amino acids (EAAs), were tested before and then during microdialysis of substance P (SP) or neurokinin A (NKA) into the dorsal horn of anesthetized monkeys. Neither peptide had significant effects on the background activity or the responses to mechanical or thermal stimulation of the skin. However, each peptide produced significant increases in the responses to iontophoretic application of one or more EAAs. In addition, following combined application of the EAAs and either SP or NKA, the responses of the cells to mechanical stimulation of the skin increased. Combined application of SP and NKA failed to produce an increase in responses to either the EAAs or to cutaneous stimuli that was greater than that produced by either peptide alone. It is concluded that SP and NKA produce an increase in the responses of STT cells to iontophoretic applications of EAAs and the combined effects of these compounds produce sustained increases in responses to mechanical stimulation of the skin. These changes mimic those observed when STT cells are sensitized by peripheral noxious stimuli, suggesting that the mechanism of induction and expression of sensitization involves the facilitation of dorsal horn neuron responses to EAAs by tachykinins.

Animals↗

Inhibition of primate spinothalamic tract neurons by stimulation in periaqueductal gray or adjacent midbrain reticular formation.

Recordings were made from spinothalamic tract (STT) cells in the lumbosacral enlargement of anesthetized monkeys. The cells were identified by antidromic activation from the contralateral ventral posterior lateral nucleus of the thalamus. Electrical stimulation at sites within the periaqueductal gray, the adjacent midbrain reticular formation, or the deep layers of the tectum were found to inhibit the activity of STT cells. In general, midbrain stimulation inhibited the background discharges and the responses of wide dynamic range cells evoked by innocuous and noxious cutaneous stimulation (29 of 37 cases). However, for six cells, midbrain stimulation preferentially inhibited the responses to noxious stimulation. The evoked responses of all 10 high-threshold cells were inhibited. In only two cases was midbrain stimulation ineffective, and no excitatory effects were observed. The mean latency to onset of inhibition resulting from midbrain stimulation was 24.9 +/- 7.2 ms (n = 35). The amount of inhibition produced by midbrain stimulation was graded with stimulus intensity. For example, trains of stimuli (333 Hz) at 50 microA produced a mean inhibition to 81.7 +/- 16.6% of control, while 200 microA resulted in a mean inhibition to 36.3 +/- 21.7%. Not only was the inhibition increased by the use of stronger current intensities, but the duration of inhibition was prolonged. Midbrain stimulation inhibited the responses of STT cells to volleys in both the A-fibers and the C-fibers of the sural nerve. However, there was a selective action in that the responses to C-fiber volleys were more strongly inhibited than were the responses to A-fiber volleys. Lesions placed in the white matter of the upper cervical spinal cord reduced the inhibition produced by stimulation in either the midbrain or the nucleus raphe magnus. The extent to which the inhibition was reduced was proportional to the extent of the cord lesions. However, even when there was an interruption of the entire lateral funiculus on the side of an STT cell and of the dorsal quadrant of the contralateral side, there was still substantial inhibition following stimulation in either brain stem site. It is concluded that while part of the inhibition is mediated by pathways descending in the dorsal lateral funiculus (DLF), at least some depends on pathways coursing through the ventral spinal cord. Inhibition of STT cells may contribute to the neuronal mechanism of the analgesia that results from stimulation in the periaqueductal gray matter in awake, behaving animals.

Animals↗

Nitric oxide-mediated spinal disinhibition contributes to the sensitization of primate spinothalamic tract neurons.

This study concentrated on whether an increase in spinal nitric oxide (NO) diminishes inhibition of spinothalamic tract (STT) cells induced by activating the periaqueductal gray (PAG) or spinal glycinergic and GABAergic receptors, thus contributing to the sensitization of STT neurons. A reduction in inhibition of the responses to cutaneous mechanical stimuli induced by PAG stimulation was seen in wide dynamic range (WDR) STT cells located in the deep layers of the dorsal horn when these neurons were sensitized during administration of a NO donor, 3-morpholinosydnonimine (SIN-1), into the dorsal horn by microdialysis. In contrast, PAG-induced inhibition of the responses of high-threshold (HT) and superficial WDR STT cells was not significantly changed by spinal infusion of SIN-1. A reduction in PAG inhibition when STT cells were sensitized after intradermal injection of capsaicin could be nearly completely blocked by pretreatment of the dorsal horn with a NO synthase inhibitor, 7-nitroindazole. Moreover, spinal inhibition of nociceptive activity of deep WDR STT neurons elicited by iontophoretic release of glycine and GABA agonists was attenuated by administration of SIN-1. This change paralleled the change in PAG-induced inhibition. However, the inhibition of HT and superficial WDR cells induced by glycine and GABA release did not show a significant change when SIN-1 was administered spinally. Combined with our recent results, these data show that the effectiveness of spinal inhibition can be reduced by the NO/cGMP pathway. Thus disinhibition may constitute one mechanism underlying central sensitization.

Animals↗

Analysis of the perception of and reactivity to pain and heat in patients with wallenberg syndrome and severe spinothalamic tract dysfunction.

BACKGROUND: The aim of the study was to assess the consequences of severe spinothalamic tract lesions resulting from lateral medullary infarct and to show that a specific pain perception can be elicited by strong thermal stimulation. CASE DESCRIPTIONS: Both patients examined presented with severe thermoalgic dissociation of the limbs contralateral to the lesion, with normal discriminative somatosensory perception and motor strength. They reported pain perception when touching very warm (>50 degrees C to 60 degrees C) objects and a brisk, occasionally uncontrolled withdrawal reaction of the arm and hand under the same conditions, without any perception of the heat nature of the stimulus. Warm stimulation, <45 degrees C, elicited no thermal perception or discrimination. Pain perception could be elicited in both patients by increasing the temperature, with a reproducible threshold of 47 degrees C to 49 degrees C. Pain always occurred after a prolonged delay of 8 to 10 seconds in response to threshold heat, and was described as deep and osseous, and clearly different from that perceived on the nonaffected side. The delay was much shorter when the temperature was increased by 4 degrees C to 5 degrees C. Cold stimulation elicited similar pain perception in one patient. Analysis of subjective perception of laser stimulation showed a much higher pain threshold on the affected hand. There were no laser-evoked potentials on this side, which suggested major spinothalamic injury. Assessment of the RIII noxious reflex revealed persistent response withdrawal reactions, with an increased threshold on the affected side, and partial consciousness of the noxious nature of the stimulus. CONCLUSIONS: To our knowledge, this is the first description of the appearance of pain perception of high temperatures in patients with severe spinothalamic injury who are suffering from a complete loss of temperature perception. This implies that noxious thermal stimulation can still be perceived via extra spinothalamic pathways (which are slow and multisynaptic), such as the spinoreticulothalamic tract. Patients with Wallenberg syndrome should be informed and made aware of their residual perception of and reactions to noxious stimulation.

Central Nervous System Diseases↗

Significant differences in the retrograde labeling of spinothalamic tract cells by horseradish peroxidase and the fluorescent tracers fast blue and diamidino yellow.

The laminar distributions of spinothalamic tract cells retrogradely labeled by the fluorescent tracers Fast Blue and Diamidino Yellow and by free or lectin-coupled horseradish peroxidase have been found to be significantly different. The total numbers of cells labeled by each method are similar, but nearly twice as many lamina I cells are labeled by the fluorescent tracers and more lamina V cells are labeled by peroxidase. Injection site spread and spurious labeling due to leakage or fibers of passage do not account for these differences. These results indicate that both horseradish peroxidase and fluorescent tracers may be selectively transported and, thus, that the cautious use of both methods should be recommended for analyses of afferent populations.

Amidines↗

Responses of spinothalamic tract cells to mechanical and thermal stimulation of skin in rats with experimental peripheral neuropathy.

1. Responses of spinothalamic tract (STT) neurons to mechanical and thermal stimulation of skin were recorded under urethane and pentobarbital anesthesia in 12 control rats and in 20 rats with experimental neuropathy. Activity of the STT cells in neuropathic rats was recorded 7, 14, and 28 days after inducing the neuropathy by placing four loose ligatures on the sciatic nerve. 2. All neuropathic animals showed guarding of the injured hindpaw and a shorter withdrawal latency from a radiant heat source of the neuropathic hindpaw than that of the sham-operated paw. 3. STT neurons in neuropathic animals showed the most profound changes 7 and 14 days after the nerve ligation. When compared with STT cells in unoperated animals, approximately half of the neurons had high background activity, responses to innocuous stimuli represented a larger percentage of the total evoked activity in wide dynamic range neurons, and the occurrence and magnitude of afterdischarges to mechanical and thermal stimuli were increased. 4. The mean threshold temperatures of heat-evoked responses of the STT cells in neuropathic animals were not different than those of cells from control animals. However, in neuropathic rats, cells reacting to small heat stimuli usually already had afterdischarges. 5. The increase in the background activity of STT cells is consistent with behavioral observations of spontaneous pain in this model of experimental neuropathy. Furthermore, the afterdischarges of STT cells may parallel the prolonged paw withdrawal in response to noxious stimuli that is seen in these animals and that is evidence for hyperalgesia. However, there was no indication of a lowered threshold for thermal stimuli as might be expected if the animals have thermal allodynia. Mechanical allodynia may have resulted from a relative increase in responsiveness to innocuous mechanical stimuli. However, responses to noxious mechanical stimuli were reduced compared with control, at least at 28 days after the ligation. Peripheral and central mechanisms responsible for the changes in responses of STT cells in neuropathic animals are suggested.

Animals↗

Responses of spinothalamic tract cells in the cat cervical spinal cord to innocuous and graded noxious stimuli.

The response properties of spinothalamic tract (STT) cells in the dorsal horn of the cervical spinal cord were examined in chloralose-anesthetized cats. The activity of 56 STT cells located in laminae IV-VI was studied, with most activity isolated in the lateral part of the dorsal horn. The level of background activity in STT cells was low (mean = 1.2 impulses/sec; n = 26). Conduction velocity estimates for STT axons ranged from 9 to 76 m/sec (mean = 38 m/sec; n = 56) and were not correlated with the recording site in the spinal cord. Most cells were antidromically activated from an electrode in the medial part of the posterior group of nuclei in the thalamus. Excitatory receptive fields were ipsilateral to the recording site, and for 38 of 40 neurons were confined to the forelimb. Although receptive fields were often restricted to part of the paw, they did not include glabrous skin. Among 31 cells classified, four groups were identified: low-threshold (LT) cells (13%) responded to pressure and brushing of the skin; high-threshold (HT) cells (13%) responded only to noxious pinching or squeezing of the skin; wide-dynamic-range (WDR) cells (58%) responded to innocuous mechanical stimuli but had a greater response to noxious stimuli; deep (D) cells (16%) responded to manipulation of subcutaneous tissues such as muscle. Heat stimuli 30 sec in duration, in the range of 43-55 degrees C, were applied to the receptive fields of 14 neurons that included representatives from all three groups with cutaneous input. Nine neurons responded to heat with thresholds that ranged from 47 degrees to 55 degrees C (mean = 51 degrees C). The responses of these nine STT cells increased with increasing stimulus intensity in the noxious range. In the cat cervical dorsal horn, STT cells can signal the occurrence of noxious stimuli on the body surface, and, judging by the sizes of their peripheral receptive fields, are capable of signaling precise information about the location of the damage. Furthermore, some cells are able to signal the intensity of a noxious heating pulse.

Animals↗

A further examination of effects of cortical stimulation on primate spinothalamic tract cells.

1. Stimulation of the sensorimotor cortex was found to excite and/or inhibit nociceptive spinothalamic tract cells. Thirteen wide dynamic range cells were inhibited by cortical stimulation, 6 were excited and 14 were both excited and inhibited. Four of six high-threshold cells were excited and one was inhibited. 2. Intermediate (200 ms) or long (2 s) duration conditioning trains were effective in reducing responses of spinothalamic cells evoked by noxious mechanical or thermal stimuli and by A- and C-fiber volleys in the sural nerve. Preferential inhibition of low-threshold responses with little or no effect on high-threshold discharges was observed in some cases. 3. Inhibitory actions were obtained primarily from stimulation of the SI sensory cortex and area 5, while excitation or excitation followed by inhibition was the dominant effect from motor cortex (area 4). Spinothalamic cells were also excited by stimulation of the medullary pyramid. 4. In eight animals extensive mapping of the sensorimotor cortex showed that for a given cell, stimulation of the sensory cortex produced inhibition while stimulation of motor cortex resulted in excitation. 5. The average latency of inhibition from sensory cortex was 29.8 +/- 10 ms, while the average latency of excitation from motor cortex was significantly shorter, 13.5 +/- 9 ms. The shortest latencies for excitation from pyramidal stimulation in the cases evaluated ranged from 2 to 9 ms. 6. Spinal cord lesions were made in five animals to determine the descending pathway(s) mediating corticofugal effects. Cortical and pyramidal effects were eliminated or considerably reduced by lesions involving the dorsal part of the lateral funiculus. This observation combined with latency data suggest that the corticospinal tract may be involved in the mediation of cortical excitation, while both pyramidal and extrapyramidal pathways are likely to be involved in cortical inhibition.

Animals↗

Cerebral-evoked responses elicited by direct stimulation of the lateral spinothalamic tract in the human.

The authors recorded cerebral-evoked responses elicited by direct stimulation of the human lateral spinothalamic tract (LST) during percutaneous cordotomy to investigate central conduction of noxious stimuli. These responses consisted of four negative potentials, peak latency being 3.8 (N1), 8.4 (N2), 12.2 (N3) and 21.9 (N4) ms respectively. N1 showed wide distribution over the scalp and was considered to be of subcortical origin. N2-N4 were distributed in both the temporal and central area. The different distribution pattern of N2-N4 from conventional somatosensory-evoked potential suggested a different projection of LST from the medial lemniscus system.

Adult↗

Responses of spinothalamic tract cells in the superficial dorsal horn of the primate lumbar spinal cord.

1. The responses of thirty-five spinothalamic tract (s.t.t.) cells in or near lamina I of the dorsal horn were examined in chloralose- and barbiturate-anaesthetized monkeys (Macaca fascicularis). Many of the cells could be classified on the basis of receptive field properties as either wide dynamic range (w.d.r.) cells or as high-threshold (h.t.) cells. 2. Thalamic stimulation sites for antidromic activation of the s.t.t. cells were in or around the ventral posterior lateral nucleus. Axons of the s.t.t. cells had a mean conduction velocity of 17 m/s (33 and 14 m/s for w.d.r. and h.t. cells, respectively). Mean minimum afferent conduction velocity averaged 37 m/s (52 and 23 m/s for w.d.r. and h.t. cells, respectively). Background activity was low (mean of 2.3 impulses/s). 3. An alternative classification of the cells was based on a kappa means cluster analysis of the responses to a series of mechanical stimuli. The response profiles for a given cell were normalized, and those of the s.t.t. cells in or near lamina I were analysed along with the responses of a population of s.t.t. cells, largely in laminae IV-VI, that had been described previously. S.t.t. cells in or near lamina I were distributed amongst three of the four groups of cells determined by the cluster analysis (types 2-4). 4. Vibratory stimuli excited most of the w.d.r. but none of the h.t. cells tested. Best frequencies were 5-10 Hz (at 100 and 500 microns indentations). 5. Most w.d.r. but few h.t. cells responded to cutaneous cooling. All of the cells responded to noxious heating, but w.d.r. cells had steeper stimulus-response curves. 6. After a series of noxious heat stimuli, the thresholds for noxious heat were lowered and responses to lower-intensity noxious heat stimuli were enhanced (sensitization). However, responses to more intense stimuli were reduced (inactivation). Similar changes were seen in the responses to graded mechanical stimuli. 7. It is concluded that s.t.t. cells in or near lamina I can signal noxious cutaneous stimuli but have poor coding abilities for innocuous mechanical stimuli. Some of these cells respond to innocuous thermal stimuli, but their role in thermoreception is unclear. The small receptive fields suggest that these cells could contribute to stimulus localization.

Action Potentials↗

The role of NMDA and non-NMDA excitatory amino acid receptors in the excitation of primate spinothalamic tract neurons by mechanical, chemical, thermal, and electrical stimuli.

The role of excitatory amino acids (EAAs) in the excitation of monkey spinothalamic tract (STT) neurons following activation of cutaneous primary afferent fibers by noxious and non-noxious stimuli was investigated. The responses of STT neurons to either NMDA or non-NMDA EAA ligands were blocked by infusion of specific antagonists through a microdialysis fiber into the region surrounding the cells. Our results show that blockade of non-NMDA receptors results in a nearly complete elimination of the responses of STT neurons to all stimuli. Blockade of NMDA receptors results in an attenuation of the responses to noxious stimuli but, in addition, prevents the development of the sensitization of STT neurons that is often observed after intradermal injection of capsaicin. These observations further support a role of EAAs in the transmission of sensory information from primary afferent fibers to dorsal horn neurons and a role for NMDA receptors in the generation of hyperalgesia.

2-Amino-5-phosphonovalerate↗

Changes in the response states of primate spinothalamic tract cells caused by mechanical damage of the skin or activation of descending controls.

1. The responses of a population of 318 spinothalamic tract (STT) cells to mechanical stimulation of the skin were recorded in anesthetized macaque monkeys by several teams of investigators. The responses were subjected to k-means cluster analysis, a multivariate statistical procedure. 2. For an analysis that pertained to the responsiveness of the neurons, the mean responses to four standard mechanical stimuli (Brush, Pressure, Pinch, and Squeeze) were used. Although no true clusters were found, the cells could be partitioned into four groups (called clusters a, b, c, and d) that responded progressively more vigorously to the stimuli. 3. For an analysis that pertained to the selectivity of the cells for various stimulus intensities, from innocuous to highly noxious, the data were normalized by taking the ratio of the mean response evoked by each stimulus to the sum of the responses and multiplying by 100. This procedure does not have a bias toward selection of any particular number of clusters and resulted in three clusters of STT cells. 4. Cluster 1 STT cells responded best to Brush. Cluster 2 cells responded weakly to Brush and Pressure and maximally to Pinch. Cluster 3 cells responded weakly to Brush, Pressure, and Pinch and maximally to Squeeze. 5. The response states of STT cells with respect to mechanical stimulation of the skin can be defined by their cluster assignments on the basis of the responsiveness (clusters a-d) and selectivity (clusters 1-3) of the cells. The response states of newly recorded STT cells can be determined by discriminant analysis from the nearest centroids of the two types of clusters in the reference population of STT cells. 6. No consistent changes in response state were detected when a second series of mechanical stimuli was applied 1 cm from the site stimulated initially or when the stimulus series was alternately repeated at the initial site and at progressively more proximal sites. However, when the stimulus series was applied five times to the initial site, the response state of five of eight cells tested showed a change. Although a change in response state required repetitive damage, even a single stimulus series increased background activity and responses to Brush at undamaged sites. 7. The background activity and responses to Brush and Pressure of all five STT cells recorded in the superficial laminae increased after repeated testing. The background activity of five STT cells recorded in the nucleus proprius also increased, but the responses of only three of the cells to Brush and Pressure increased.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗