Postsynaptic Dorsal Column Neurons Research Articles

Comparative study of viscerosomatic input onto postsynaptic dorsal column and spinothalamic tract neurons in the primate.

The purpose of the present investigation was to examine, in the primate, the role of the postsynaptic dorsal column (PSDC) system and that of the spinothalamic tract (STT) in viscerosensory processing by comparing the responses of neurons in these pathways to colorectal distension (CRD). Experiments were done on four anesthetized male monkeys (Macaca fascicularis). Extracellular recordings were made from a total of 100 neurons randomly located in the L(6)-S(1) segments of the spinal cord. Most of these neurons had cutaneous receptive fields in the perineal area, on the hind limbs or on the rump. Forty-eight percent were PSDC neurons activated antidromically from the upper cervical dorsal column or the nucleus gracilis, 17% were STT neurons activated antidromically from the thalamus, and 35% were unidentified. Twenty-one PSDC neurons, located mostly near the central canal, were excited by CRD and three were inhibited. Twenty-four PSDC neurons, mostly located in the nucleus proprius, did not respond to CRD. Of the 17 STT neurons, 7 neurons were excited by CRD, 4 neurons were inhibited, and 6 neurons did not respond to CRD. Of the unidentified neurons, 23 were excited by CRD, 7 were inhibited, and 5 did not respond. The average responses of STT and PSDC neurons excited by CRD were comparable in magnitude and duration. These results suggest that the major role of the PSDC pathway in viscerosensory processing may be due to a quantitative rather than a qualitative neuronal dominance over the STT.

GABAergic Neurons that Contain Neuropeptide Y Selectively Target Cells with the Neurokinin 1 Receptor in Laminae III and IV of the Rat Spinal Cord

Neuropeptide Y (NPY) is contained in a population of GABAergic interneurons in the spinal dorsal horn and, when administered intrathecally, can produce analgesia. We previously identified a strong monosynaptic link between substance P-containing primary afferents and cells in lamina III or IV with the neurokinin 1 (NK1) receptor. Because some of these cells belong to the spinothalamic tract, they are likely to have an important role in pain mechanisms. In this study, we used confocal microscopy to examine the input to lamina III/IV NK1 receptor-immunoreactive neurons from NPY-containing axons. All of the cells studied received a dense innervation from NPY-immunoreactive axons, and electron microscopy revealed that synapses were often present at points of contact. Most NPY-immunoreactive boutons were also GABAergic, which supports the suggestion that they are derived from local neurons. The association between NPY-containing axons and NK1 receptor-immunoreactive neurons was specific, because postsynaptic dorsal column neurons (which were located in laminae III-V but did not possess NK1 receptors) and lamina I neurons with the NK1 receptor received significantly fewer contacts from NPY-immunoreactive axons. In addition, the NK1 receptor-immunoreactive lamina III/IV cells received few contacts from nitric oxide synthase-containing axons (which belong to a different population of GABAergic dorsal horn neurons). The NPY-containing axons appeared to be targeted to the NK1 receptor-immunoreactive neurons themselves rather than to their associated substance P-immunoreactive inputs. The dense innervation of these cells by NPY-containing axons suggests that they may possess receptors for NPY and that activation of these receptors may contribute to NPY-mediated analgesia.

Electrophysiological evidence that spinomesencephalic neurons in the cat may be excited via spinocervical tract collaterals

Extracellular microelectrode recordings were made from spinomesencephalic tract (SMT) neurons in the lumbosacral spinal cord of cats anaesthetized with chloralose and paralysed with gallamine triethiodide. The SMT cells were antidromically fired from the posterolateral parts of the superior colliculus and the intercollicular region, were located in laminae IV to VIII, and had response properties and axonal conduction velocities similar to those described previously. The effects of stimulating the dorsolateral funiculus of the cervical cord at C3 and rostral C1, below and above the termination of spinocervical tract (SCT) axons in the lateral cervical nucleus, were examined on 33 SMT cells. The strength of stimulation was adjusted so that at C3 it was above threshold for antidromic activation of SCT cells and at C1 was below threshold for activation of the same cells. Seven (21%) SMT neurons were excited from C3 but not from C1. The remaining 26 (79%) were excited from both C3 and rostral C1 and 23 (70% of these) were excited significantly more from C3. That is, 91% of the total sample were either excited only from C3 or more strongly from C3 than from rostral C1. We discuss the possible neuronal systems involved and conclude that the greater excitatory effects from C3 are most likely due to antidromic activation of the SCT. The shortest latency effects from C3 indicate a monosynaptic linkage between SCT cells with the fastest axons and the SMT. The longer latency actions may be due to monosynaptic connexions from SCT cells with slower conducting axons, to di- or polysynaptic actions from SCT cells with fast axons, or a combination of both. SMT cells are another population of spinal neurons, in addition to postsynaptic dorsal column, spinothalamic and dorsal horn spinocerebellar neurons, which receive excitation via SCT collaterals.

Pelvic visceral input into the nucleus gracilis is largely mediated by the postsynaptic dorsal column pathway.

1. The purpose of this study was to investigate a proposed role for the postsynaptic dorsal column (PSDC) pathway in mediating visceral nociceptive input into the dorsal column (DC) nuclei. 2. In one group of animals, the hypogastric nerves were sectioned, thereby restricting colorectal input into the cord to pelvic afferent pathways known to coverage on lower lumbar and sacral segments. Extracellular recording were made from 41 nucleus gracilis (NG) cells that responded to colorectal distension (CRD). Results reported are from 15 NG cells that were tested before and after the administration of morphine into the sacral cord by microdialysis. 3. The responses of 11 NG cells to CRD were dramatically reduced by morphine infused into the sacral cord through a microdialysis fiber. This reduction was reversed by an intravenous injection of naloxone. Microdialysis administration of 6-cyano-7-nitro-quinoxaline-2,3-dione (CNQX) or a lesion of the DC also abolished the responses of the NG cells to CRD. 4. Four NG cells that responded to CRD showed an increase in their background activity approximately 25 min after an injection of mustard oil (MO). This increase in activity was counteracted by morphine or by a lesion of the DC. 5. In a second group of animals, recordings were made from 28 PSDC cells in the L0-S1 segments of the cord. These units were antidromically activated by stimulation of the upper cervical fasciculus gracilis. The projections of five PSDC neurons into the NG were traced with the use of antidromic mapping. Results are reported for the responses of 12 PSDC cells to CRD and to cutaneous stimuli before and after morphine administration into the sacral cord by microdialysis. 6. Morphine given spinally reduced the responses of 12 PSDC cells to CRD. This reduction was reversed by an intravenous injection of naloxone. CNQX administered spinally also abolished the responses to CRD of the PSDC cells tested. 7. Four other PSDC cells were studied before and after an injection of MO into the colon. Their background activity started to increase within 25 min after the injection. Morphine suppressed this increase in background activity and this effect of morphine was reversed by naloxone. 8. The responses of NG cells to cutaneous stimuli were not significantly affected by morphine in the dose used. On the other hand, morphine significantly reduced the responses of PSDC cells to noxious cutaneous stimuli although this effect was not as dramatic as that on responses to visceral stimuli. 9. From the results of the studies described in this and the companion paper, we conclude that there is an important pelvic visceral nociceptive pathway involving PSDC neurons that synapse in the NG. The NG in turn activates neurons in the ventral posterolateral (VPL) nucleus of the thalamus. We presume that activation of VPL neurons by noxious visceral stimulation contributes to visceral pain sensation and thus that pelvic visceral pain depends largely on activity in the DC-medial lemniscus system.

Colocalization of glycine and GABA in synapses on spinomedullary neurons

Spinomedullary neurons of the postsynaptic dorsal column pathway in adult cats were retrogradely labelled with horseradish peroxidase. Postembedding immunogold reactions were performed with antisera which recognise GABA or glycine to determine if synaptic boutons in contact with these neurons contain both transmitters. Analysis of series of ultrathin sections revealed that synaptic profiles with strong immunogold reactions for GABA usually also displayed strong immunogold reactions for glycine. Pre-embedding immunocytochemistry was performed on sections containing labelled cells with a monoclonal antibody which recognises the glycine receptor-associated protein, gephyrin. Many synapses onto postsynaptic dorsal column neurons were associated with gephyrin-like immunoreactivity and these typically contained irregularly shaped vesicles. Immunogold reactions showed that synaptic profiles apposed to gephyrin-immunoreactive junctions contained GABA and glycine. The evidence suggests that glycine is a neurotransmitter at synapses on spinomedullary neurons and that it is colocalized with GABA.

Synaptic organization of excitatory and inhibitory boutons associated with spinal neurons which project through the dorsal columns of the cat

The cell bodies and proximal dendrites of postsynaptic dorsal column neurons were examined for synaptic boutons which displayed immunoreactivity for the principal excitatory and inhibitory neurotransmitters, glutamate and GABA. The neurons were labelled by retrograde transport of horseradish peroxidase and GABA or glutamate-containing boutons were revealed by performing postembedding immunogold reactions on electron microscope sections. Five neurons were examined and all of them were postsynaptic to boutons which contained either GABA or glutamate. Quantitative analysis of two of the cells revealed that more than 90% of the synaptic profiles associated with them displayed immunogold reactions for these transmitters. Analysis of series of alternate sections, which were reacted for either GABA or glutamate, showed that there was no overlap in the populations of immunoreactive boutons. Furthermore, GABA and glutamate immunoreactions were associated with boutons which had different morphological characteristics. In addition, some large glutamate-enriched boutons were postsynaptic to small boutons which displayed immunogold reactions for GABA. This study demonstrates morphological bases for direct excitation, postsynaptic inhibition and presynaptic inhibition of postsynaptic dorsal column cells.

Ascending tract neurones processing information from group II muscle afferents in sacral segments of the feline spinal cord.

1. Ascending tract neurones located in the dorsal horn of sacral segments of the spinal cord have been investigated by extracellular and intracellular recording in the anaesthetized cat. The aim was to determine whether information from group II afferents that terminate within the sacral segments is conveyed to supraspinal structures and which types of neurones are involved. 2. A considerable proportion of ascending tract neurones found in the dorsal horn in the same segments as the pudendal (Onuf's) motor nucleus were excited by group II muscle afferents. The great majority (93%) of these neurones had axons ascending in ipsilateral funiculi. Spinocervical tract neurones constituted the largest proportion (82%) of such neurones, while very few spinocerebellar tract and propriospinal neurones and no postsynaptic dorsal column neurones were found among them. 3. In addition to activation by group II muscle afferents all of the neurones were strongly excited by cutaneous afferents. The most potent excitation was evoked by afferents of the posterior biceps-semitendinosus and gastrocnemius muscle nerves and by afferents of the cutaneous femoris, sural and pudendal nerves. The latencies of intracellularly recorded excitatory potentials were indicative of a high incidence of monosynaptic coupling between the afferents and ascending tract neurones. 4. The highly effective monosynaptic excitation of spinocervical tract neurones in the sacral segments by group II afferents is in contrast to the weak disynaptically mediated actions of group II afferents on such neurones in the L6-L7 segments but comparable to the actions of group II afferents on ascending tract neurones in the midlumbar segments. 5. Both the patterns of peripheral input and the latencies of synaptic actions in ascending tract neurones were similar to those in interneurones at the same locations (accompanying report). Similar information is therefore likely to be processed by both categories of neurones. 6. The role of sacral spinocervical tract neurones as a system for transmitting information from group II muscle afferents to supraspinal centres and the potential contribution of this system to the perception of limb position are discussed.

Lumbosacral spinal neurons in the cat that are candidates for being activated by collaterals from the spinocervical tract

Lumbosacral spinal neurons activated via the spinocervical tract were stained by intracellular injection of horseradish peroxidase in cats anaesthetized with chloralose and paralysed with gallamine triethiodide. The neurons were activated orthodromically by single shock stimulation of the ipsilateral dorsolateral funiculus at the second to third cervical segment, but not from the rostral part of the first cervical segment. Twenty nine cells were recovered from the histological material and subsequently reconstructed from transverse sections. Sixteen cells (55%) had axons that projected ipsilaterally to the lateral funiculus and their somata were located in two regions of the spinal cord, one group in the dorsal horn (laminae IV-V) and the other in the intermediate gray matter (laminae VI-VII). The axons of 10 of these cells gave off collaterals, and in seven of them the collaterals ramified in the grey matter deep to the cell body. The axons of five cells (17%) projected medially towards the central canal, four crossing the mid line in the ventral white commissure and ascending in the contralateral ventral funiculus. Only one of these cells had an axon collateral that crossed into the contralateral dorsal horn. Of the remaining eight cells, three had no obvious long axons but had many local axon collaterals, the axons of three cells were not stained, one had an axon projecting towards the ipsilateral ventral funiculus and one was a motoneuron and its axon projected into a ventral root. A feature of the dendritic trees of many cells was their wide spread in the mediolateral and/or the dorsoventral directions, although no dendrites reached dorsally into lamina II. Twenty-two cells (76%) were excited by moving hairs and by noxious pinch, three (10%) by hair movement alone, two (7%) by noxious pinch and pressure, and for two cells (7%) no receptive field could be found. It is concluded that not only postsynaptic dorsal column neurons receive input from the spinocervical tract but also other cells in the dorsal and ventral horns and the intermediate gray matter. Possible identities for these cells are discussed.

Organization of the serotonergic innervation of spinal neurons in rats—I. Neuropeptide coexistence in varicosities innervating some spinothalamic tract neurons but not in those innervating postsynaptic dorsal column neurons

Previous studies have suggested that peptides such as substance P and thyrotropin-releasing hormone coexist with serotonin in the same varicosities in the ventral horn and intermediate gray of the spinal cord in rat. However, coexistence of these peptides with serotonin is rare in fibers in the superficial dorsal horn. Since it has been proposed that serotonergic fibers in the superficial dorsal horn act to modulate nociception, it was hypothesized that the serotonergic neurons that contain neither substance P nor thyrotropin-releasing hormone might constitute a specifically antinociceptive subset of serotonergic neurons. This being the case, it would be expected that different types of serotonergic neurons innervate nociceptive and non-nociceptive spinal neurons. In order to test this hypothesis, a group of cells that include nociceptive neurons (spinothalamic tract neurons) and a group of predominantly non-nociceptive neurons (postsynaptic dorsal column neurons) in the spinal cord of rat were retrogradely labeled. Sections of the spinal cord containing retrogradely labeled spinothalamic tract or postsynaptic dorsal column neurons were stained for serotonin and either substance P or thyrotropin-releasing hormone using two-color immunohistochemistry. A retrogradely labeled cell was classified as “apposed” if there was no discernible distance between an immunohistochemically labeled varicosity and the cell. Eighty per cent of spinothalamic tract and 83% of postsynaptic dorsal column profiles were apposed by serotonin-immunoreactive varicosities in the spinal cord. Thirty-one per cent of the spinothalamic tract profiles that were apposed by serotonergic varicosities were apposed by serotonergic varicosities that were also stained for thyrotropin-releasing hormone. The distribution of the latter spinothalamic neurons was similar to that reported for spinothalamic tract neurons responsive to joint movement. In addition, at least 63% of the spinothalamic tract profiles which were apposed by serotonergic varicosities were apposed by “serotonin-only” varicosities, including most spinothalamic tract neurons in the marginal zone, suggesting that at least some “serotonin-only” neurons are antinociceptive. However, contrary to the hypothesis, at least 94% of the postsynaptic dorsal column profiles apposed by serotonergic varicosities were apposed by “serotonin-only” varicosities. These findings suggest that there may be a relationship between the sensory modality to which a spinal neuron responds and the type of serotonergic innervation it receives. However, it appears that “serotonin-only” neurons may not constitute a specifically antinociceptive category of serotonergic neurons.

Postsynaptic dorsal column neurons in the cat: a study with retrograde transport of horseradish peroxidase.

The numbers, laminar position, perikaryal and dendritic morphology, and axonal trajectories of postsynaptic cells ascending the dorsal column have been studied after implantation of HRP pellets in either the dorsal columns or dorsal column nuclei after destruction of the dorsolateral fascicle on one side. Observations made throughout the spinal cord gave estimated figures of 800-1000 and 1700-2000 cells in lumbosarcal and brachial enlargements respectively on the side of the implant. The commonest type (C), centred on lamina IV, had dendritic trees greatly extended rostrocaudally and restricted mediolaterally in the lateral dorsal horn, the extension and restriction diminishing for more medial cells. Type B cells differed dramatically, with large straight dendrites in the transverse plane and large perikarya in medial lamina V. Type A cells, distinguished by both rostrocaudal and mediolateral restriction in dendritic trees, were only found medially in laminae III and IV. Outside the enlargements, in high lumbar and thoracic cord, many fewer cells were found, corresponding to Type C but with dendrites much elongated rostrocaudally and little mediolateral variation. Many small fusiform cells were found in medial lamina VI in the upper cervical cord, distinct from any of the above. A few cells were found in the cord enlargements in lamina VII of the contralateral ventral horn, with axons crossing through the ventral commissure. The axons of all cell types were tortuous, and some entered the dorsolateral fascicle before crossing into the dorsal column: collaterals were often seen but could not be followed far. A complementary study of cells with axons ascending in the dorsolateral fascicle is reported in the following paper.

Descending influences on the cutaneous receptive fields of postsynaptic dorsal column neurones in the cat.

1. The influence of activity in descending systems on the cutaneous receptive field properties of postsynaptic dorsal column (PSDC) neurones has been investigated in chloralose-anaesthetized cats. The main aim of the study was to determine whether the receptive field boundaries of PSDC neurones are under the control of systems descending from the brain. 2. Single-unit recordings were made from the ascending axons of PSDC units in the dorsal columns. Receptive fields were analysed using light tactile and noxious mechanical and thermal stimuli, both before and during a reversible block of spinal conduction produced by cooling the cord rostral of the recording site. 3. The light tactile excitatory fields of PSDC neurones were largely unaffected by the cold-block procedure. 4. In contrast, both the sensitivity of PSDC neurones to noxious stimuli and the area of skin from which they could be effectively excited by such stimuli were found to be profoundly modified by interruption of descending activity. Two-thirds of the units excited by noxious pinch responded more vigorously in the cold-blocked state and one-half from an expanded area of skin. Responses to noxious radiant heat were similarly modified. 5. Inhibition evoked in PSDC neurones, whether by light tactile or noxious stimuli, involved predominantly segmental mechanisms since it remained effective in the cold-blocked state. 6. It is concluded that neurones of the PSDC system are amongst those dorsal horn neurones with receptive field geometries which may be modified by activity in descending systems.

Spinocervical tract-dorsal column postsynaptic neurons: a double-projection neuronal system.

Evidence is presented for the existence of a newly discovered double-projection spinal neuronal system, the spinocervical tract-dorsal column postsynaptic neurons. The neurons are characterized by axonal bifurcation in the cervico-thoracic junction, by branched axons traveling in the dorsal column and the dorsolateral funiculus, and by double projection to the dorsal column nuclei and the lateral cervical nucleus. The neurons, with longitudinally distributed dendritic trees and local axon collaterals, primarily originate in laminae III-V of the dorsal horn and receive innocuous and noxious inputs transmitted along the A-beta primary afferents from the periphery. These neurons are thought to be an intersection of the spinocervical tract and the dorsal column postsynaptic neurons, and to function as a nonlemniscal system in mediation and modulation of ascending sensory information, including pain.

Temporally dependent changes in response properties of dorsal horn neurons after dorsolateral funiculus lesions

Previous studies in this laboratory have shown 1) that 19% of L6 and L7 dorsal horn cells in normal cats respond only with excitatory postsynaptic potentials (EPSPs) to sural nerve stimulation, and 2) that the distribution of dorsal horn neurons responding with impulses to sural nerve stimulation is increased in cats with chronic lateral funiculus lesions. The present study was undertaken to determine whether strengthening of subliminal sural nerve projections could account for the changes seen after lateral funiculus lesions and to explore the nature of these changes in greater detail. Ipsilateral L6 and L7 dorsal horn cells of cats with T12 dorsolateral funiculus (DLF) lesions were studied electrophysiologically at less than 1-30 days postoperatively (DPO) and were compared with similar cells recorded in normal cats. The major results were as follows. 1. Responsiveness to peripheral stimulation was depressed for up to 3 DPO following the lesions. 2. The percentage of L6 and L7 dorsal horn cells showing spontaneous activity was elevated at 3 DPO and declined to normal levels by 28 DPO. 3. The percentage of cells that responded to sural nerve stimulation increased over 3-30 DPO and was significantly greater than normal at 28-30 DPO. The increase in the percentage of cells giving impulses to sural nerve stimulation at 28-30 DPO was similar to the percentage of cells with subliminal responses to sural nerve stimulation in normal cats. A subpopulation of identified postsynaptic dorsal column neurons also showed a significant increase in the percentage that responded to sural nerve stimulation after DLF lesions. 4. The percentage of cells that had a cutaneous receptive field (RF) overlapping the region of skin innervated by the sural nerve also increased over time and was significantly greater than normal at 28-30 DPO. 5. Median low-threshold or high-threshold RF areas were not significantly greater than normal ipsilateral to DLF lesions at any survival time. 6. The proportions of low-threshold (LT), high-threshold (HT), and multireceptive (MR) cells were not significantly different from normal at any survival time from less than 1 to 30 DPO.(ABSTRACT TRUNCATED AT 400 WORDS)

Cutaneous excitatory and inhibitory input to neurones of the postsynaptic dorsal column system in the cat.

1. In chloralose-anaesthetized cats single-unit microelectrode recordings were made from axons in the dorsal columns, at the lumbar level, identified as belonging to the postsynaptic dorsal column (PSDC) system. 2. Excitatory and inhibitory receptive field arrangements of a sample of seventy-five PSDC neurones were examined in detail using natural cutaneous stimuli. 3. The sample was characterized by a high degree of convergent input: 80% of units were activated by both light tactile and noxious mechanical stimuli and more than half of those examined were excited by noxious radiant heat. In addition, three-quarters of the units had inhibitory receptive fields on the ipsilateral limb. 4. Twenty-three units (27%) were influenced by input from areas of both hairy and glabrous skin covering the foot and distal limb. Neurones in this group had complex receptive fields, many of which occupied several discontinuous areas of skin. Background and evoked activity of these units could frequently be inhibited by light tactile and/or noxious stimuli. Their inhibitory receptive fields occupied small areas of skin overlapping or adjacent to excitatory fields. 5. Fifty-two units (73%) had receptive fields restricted to areas of hairy skin on the thigh and upper hindlimb. Half the units in this group had coextensive low- and high-threshold excitatory areas but about one-third had a concentric receptive field organization; a high-threshold excitatory component extending beyond, or around, a central low-threshold area. The discharge of these units could be inhibited only by light tactile stimuli. Their inhibitory receptive fields covered extensive areas of skin, sometimes completely surrounding the excitatory field. 6. The complex receptive field arrangements observed for neurones of the postsynaptic dorsal column system are discussed in relation to previous observations on dorsal horn neurones of other ascending tracts.

Fine structure of synapses associated with characterized postsynaptic dorsal column neurons in the cat

Fourteen dorsal horn neurons with axons projecting through the dorsal columns were identified either by electrophysiological methods (and subsequently injected with horseradish peroxidase) or by retrograde labelling with horseradish peroxidase in cats. All neurons were contacted by small (<2 μm) boutons containing spherical or elongated agranular vesicles. One neuron with its soma located in lamina III received additional contacts from central elements of glomerular complexes. Neurons with somata located more ventrally (deep lamina IV and V) were also postsynaptic to large (>2 μm) electron lucent profiles which formed multiple synapses with the labelled cells. Some boutons presynaptic to postsynaptic dorsal column neurons were themselves postsynaptic to profiles containing pleiomorphic agranular vesicles at axoaxonic synapses. They also occasionally participated in triadic complexes. It is concluded that the synaptic arrangements formed by boutons in association with postsynaptic dorsal column neurons differ significantly from those associated with spinocervical neurons. Such differences might provide the anatomical substrate for the observed receptive field characteristics of these neurons.

Electrical stimulation reveals relatively ineffective sural nerve projections to dorsal horn neurons in the cat

Electrical stimulation of the sural nerve (SN) revealed input from sural nerve afferents to L6 and L7 dorsal horn neurons that were not apparent using natural mechanical stimuli, especially in cells with variable latency responses to SN stimulation. Nearly all (31/32) cells that had reliable, fixed latency responses to SN stimulation also had an excitatory receptive field (RF) in the region of skin innervated by the sural nerve (SN region). About one-third (20/57) of the cells with variable latency responses to SN stimulation, however, had an RF outside the SN region. Most (130/146) cells with no response to SN stimulation had RFs outside the SN region. There were no obvious differences between variable latency cells with RFs in the SN region vs those with RFs outside it in latency of response to SN stimulation, recording depth, RF sizes or modality properties. In a subsample of 31 postsynaptic dorsal column neurons all cells responding to SN stimulation also had an RF in the SN region. Strengthening of relatively ineffective projections from the sural nerve by lesions might be expected to lead to an increase in the proporpotion of cells responding with impulses to natural stimulation of the skin innervated by the sural nerve, and, hence, to an increase in average RF size.

Dorsal column postsynaptic neurons in the cat are excited by myelinated nociceptors

Some (25-50%) dorsal column postsynaptic (DCPS) neurons respond only to innocuous mechanical stimuli; the remainder (50-75%) responds to both innocuous and noxious mechanical stimuli. Those that respond to noxious mechanical stimuli (pinch) are assumed to be excited by input from nociceptive primary afferents, but it is conceivable that their pinch-evoked responses are produced by the inadvertent activation of those low-threshold mechanoreceptive primary afferents that respond to stretching the skin. Because nociceptive primary afferents respond reliably to noxious heat and low-threshold mechanoreceptors do not, we tested DCPS neurons in the cat lumbar spinal cord with a series of noxious heat stimuli (48 degrees C or 50 degrees C-56 degrees C; 30 s duration). Seven of eight pinch-responsive neurons responded to noxious heat, but only after their receptive fields had been sensitized by prolonged or repeated heating. The results show that (1) many DCPS neurons in the cat are excited by nociceptive primary afferents and (2) these nociceptive afferents are probably myelinated high-threshold mechanoreceptors.

Postsynaptic dorsal column pathway of the rat. II. Evidence against an important role in nociception

The response characteristics of neurons at the origin of the postsynaptic dorsal column (PSDC) pathway were determined in unanesthetized, decerebrated, spinalized rats. Sixty-four percent of PSDC neurons responded only to innocuous mechanical stimuli. Thirty-six percent responded to innocuous stimuli but were more powerfully activated by noxious pinch, Ninety-three percent of the tested PSDC neurons were not activated by any of several intensities of sustained, repeated noxious heating of their receptive fields. The failure of pinch-responssive PSDC cells to respond to thermal stimulation, even in sensitized skin, suggests that they do not receive a functionally significant input from C polymodal nociceptors, heat nociceptors, or mechanical-heat nociceptors. We conclude, therefore that the postsynaptic dorsal column pathway is not importantly involved in nociception in the rat.

Effects of hind limb nerve section on lumbosacral dorsal horn neurones in the cat.

The sciatic and saphenous nerves of one hind limb were sectioned in young adult cats anaesthetized with halothane. Between 19 and 55 days later, under chloralose anaesthesia, dorsal horn neurones in the L6 and L7 segments were recorded and their receptive field properties examined. In seven animals recordings were made from identified spinocervical tract, post-synaptic dorsal column and dorsolateral funicular neurones as well as from neurones that did not project through these pathways. Thirty-one neurones were intracellularly stained with horseradish peroxidase, and fifty-three were recorded extracellularly and located by reference to stained cells. In two animals (both 31 days after nerve section) no attempt was made to identify axonal projections of the dorsal horn neurones in order to avoid any effects of cervical cord search stimuli on the cells' properties, but all isolated extracellularly recorded units were examined. On the side ipsilateral to the nerve sections 143 units were recorded. In all experiments, neurones in the medial three-quarters of the dorsal horn had no discernible cutaneous, mechanosensitive receptive fields between 19 and 55 days after nerve section. There were only two exceptions to this generalization, one neurone being one of the most rostral cells in the sample (in caudal L5) and the other being one of the most caudal cells (in caudal L7). We present evidence to show that neither of these two neurones had inappropriate receptive fields in terms of the somatotopic organization of the dorsal horn. All other neurones with receptive fields on the skin were appropriately located in the somatotopic map laid out in the dorsal horn. There was no evidence for gross anatomical changes in the dendritic trees of dorsal horn neurones following sciatic and saphenous nerve sections. We have been unable to confirm that, following loss of cutaneous receptive fields by peripheral nerve section, dorsal horn neurones in adult cats acquire 'inappropriate' receptive fields. Possible reasons for this are discussed.

The morphology of dorsal column postsynaptic spinomedullary neurons in the cat.

Dorsal column postsynaptic ( DCPS ) spinomedullary neurons from the cat's lumbosacral enlargement were identified by antidromic stimulation of the cervical dorsal columns and stained intracellularly with horseradish peroxidase. The cell bodies were located in laminae III-IV. Their dendritic arbors were elongated rostrocaudally but narrow mediolaterally. On the average, the arbors were X 5 longer than they were wide. Most of the neurons had nearly all of their dendrites in laminae III-IV and some of the neurons had, in addition, a considerable amount of dendritic surface area in lamina V. Only one neuron had more than a very small amount of dendritic surface area dorsal to lamina III. Seven of the neurons had unmyelinated axon collaterals that arborized extensively and issued varicosity-bearing terminal branches in laminae III-V, both within and beneath their dendritic territories. All of the neurons were excited by myelinated, low-threshold mechanoreceptors. Since the rostrocaudally elongated and mediolaterally narrow dendritic arbors of DCPS neurons are in register with the laminae III-IV terminal distributions of myelinated, low-threshold mechanoreceptors, it is probable that this excitation arises from a monosynaptic and topographically discrete innervation. About one-half of the DCPS neurons were also excited by noxious stimuli. It is probable that this excitation is accomplished by a polysynaptic pathway since DCPS dendritic arbors and nociceptor terminal distributions are largely or completely separate.