Lissauer's Tract Research Articles

Morphology and distribution of primary afferent fibres expressing alpha-galactose extended oligosaccharides in the spinal cord and brainstem of the rat. Light microscopy.

The light microscopic morphology and distribution of non-substance P-containing small primary afferent fibres were studied. These fibres were labelled using LD2 and LA4 monoclonal antibodies which recognize alpha-galactose extended oligosaccharides expressed by primary afferent neurons. The LD2 and LA4 antibodies immunostained small primary afferent fibres ending mainly in lamina II of the spinal cord dorsal horn and trigeminal subnucleus caudalis of the rat. The lamination pattern of both types of primary afferents was assessed using an image analysis system. The highest density of LD2-immunoreactive fibres was located in a patchy band located in lamina II outer, while LA4-immunoreactive fibres were distributed mainly through lamina II inner. In lateral regions of cervical and lumbar dorsal horn the LA4-immunoreactive band is broader and comprises almost all lamina II. In contrast to substance P-containing primary afferents, a low density of LD2- or LA4-immunoreactive fibres was found in lamina I, and no terminal fields were found in lamina V or lamina X of the spinal cord or in levels of the trigeminal system outside the subnucleus caudalis. Both antibodies also labelled the parent fibres in the white matter fasciles. LD2-immunoreactive fibres were located in the dorsal roots, medial regions of the Lissauer tract, dorsal columns of the spinal cord, outer regions of the spinal trigeminal tract and dorsal to the cuneatus and gracilis nuclei. In contrast, LA4-immunoreactive fibres were restricted to the dorsal roots, medial and lateral regions of the Lissauer tract and the outer regions of the trigeminal tract. Immunostained fibres in the rootlets of the X and IX nerves and immunoreactive terminal arborizations in various subnuclei of the nucleus tractus solitarius were seen using both antibodies. These results show that subpopulations of small primary afferents stained by LD2 and LA4 antibodies have distinct patterns of central distribution and are consistent with a subdivision of small primary afferents into peptide- and non-peptide-containing groups.

Calcitonin gene-related peptide immunoreactivity in the cat lumbosacral spinal cord and the effects of multiple dorsal rhizotomies.

This study determined the extent of the rostral projection of calcitonin gene-related peptide-like immunoreactive (CGRP-IR) primary afferents in the cat lumbosacral spinal cord. To do this we examined the distribution of CGRP-like immunoreactivity (CGRP-LI) contralateral and ipsilateral to multiple dorsal rhizotomies. In the contralateral dorsal spinal cord, CGRP-IR fibers were mostly observed in Lissauer's tract, the dorsal columns, and laminae I, II, and V. Fewer CGRP-IR fibers were observed in laminae III, IV, and VI and the area around the central canal. The location of the CGRP-LI suggests that the afferents arose from nociceptors. Unilateral dorsal rhizotomies of five consecutive segments in the lumbar enlargement caused a substantial although incomplete loss of CGRP-LI in the rhizotomized dorsal spinal cord ipsilateral to the lesions. The majority of the remaining CGRP-IR fibers were located in Lissauer's tract, the dorsal columns, and the lateral part of laminae I and V. Ventral rhizotomies or an ipsilateral hemisection in the most rostral rhizotomized segment, in addition to the dorsal rhizotomies, had no noticeable effect upon the density or location of the remaining CGRP-LI. These results suggest that the majority of the CGRP-LI within the rhizotomized region of spinal cord was contained within branches of small-diameter primary afferents that entered the spinal cord through intact dorsal roots located caudal to the rhizotomized segments of spinal cord. It is concluded that CGRP-IR small-diameter primary afferents are capable of projecting at least five segments beyond their segment of entry and supplying collaterals to the superficial and deeper layers of the dorsal horn involved in the processing of nociceptive information.

Calbindin immunoreactivity in the neurons of the spinal cord and dorsal root ganglion of the domestic fowl

The distribution of the vitamin D-dependent calcium-binding protein, calbindin, was mapped in the brachial spinal cord and in the 15th dorsal root ganglion of the domestic fowl, using fluorescence immunohistochemistry. Cell somata of the dorsal root ganglion ranged in area from 200 microns 2 to 2000 microns 2. Sixteen percent of cell bodies displayed calbindin immunoreactivity. Reactivity occurred in both the small and large sensory neurons. These were randomly distributed within each ganglion. In the spinal cord, calbindin immunoreactivity was intense in Lissauer's tract, and in nerve fibres and nerve cell bodies within laminae 1 to 3 of the dorsal horn. Scattered varicose fibres were observed in laminae 4 to 7. Immunoreactivity was intense in laminae 10 where nerve fibers formed a meshwork around the central canal. Immunoreactive perikarya were occasionally observed in the outer region of lamina 10 and between laminae 8 and 9. The perikarya of the large motoneurons of lamina 9 were not reactive although they were enmeshed in calbindin-immunoreactive fibres.

Microsurgical DREZ-otomy for the treatment of spasticity and pain in the lower limbs.

The authors report on a series of 53 bedridden patients suffering from harmful spasticity in one (6) or both (47) lower limbs, who were treated with microsurgical DREZ-otomy. Surgery was performed to treat fixed abnormal postures in flexion in 49 patients and hyperextension in 3, and, additionally, to treat pain in 37 patients. Microsurgical DREZ-otomy was introduced in 1972, on the basis of anatomical studies of the human dorsal root entry zone (DREZ) showing a topographical segregation of the afferent fibers according to their size and functional destinations. It consists of a 2 mm deep microsurgical lesion directed at a 45 degrees angle in the posterolateral sulcus and penetrating the dorsal root entry zone in its ventrolateral aspect, at the level of all the rootlets considered involved in spasticity (and pain). It destroys mainly the lateral (nociceptive) and central (myotatic) afferent fibers as well as the facilitatory medial part of the Lissauer tract, while sparing most of the medial (lemniscal) fibers, the suppressor lateral part of the Lissauer tract, and more or less of the dorsal horn (DH). The postoperative results were evaluated after a mean follow-up period of 3 years and 4 months. Both spasticity and spasms were significantly decreased or suppressed in 75% and 88.2% of the patients, respectively. When present, pain was relieved without abolition of sensation in 91.6%. These benefits--combined with complementary orthopedic surgery in 23 patients--resulted in either disappearance or marked reduction of the abnormal postures in 85.3% of the patients and of articular limitations in 96.8%. Mild-to-severe complications occurred in 25 patients and precipitated or were responsible for death in 5. This is explained by the fact that the general and neurological conditions of most of the patients--especially those affected by multiple sclerosis--were precarious. MDT has, however, enabled a majority of these severely disabled patients to sit and lie comfortably, and has allowed them to reach a significantly improved quality of life.

Ultrastructural localization of neuropeptides and GABA in rat dorsal horn: a comparison of different immunogold labeling techniques.

Several immunogold techniques were used to determine the ultrastructural localization of calcitonin gene-related peptide (CGRP), tachykinin, somatostatin, and gamma-amino-butyric acid (GABA) immunoreactivity in the dorsal horn of rat spinal cord. The immunocytochemical reactions were carried out directly on ultrathin sections from non-osmicated frozen tissue, non-osmicated low temperature-embedded (Lowicryl K4M) tissue, and osmicated epoxy-embedded material. Preservation of ultrastructural morphology and immuno-labeling efficiency were compared. Morphology of subcellular organelles was relatively good in ultra-thin frozen sections, which showed the highest immunoreactivity. However, only very small samples of tissue could be examined. Although there was relatively good immunolabeling in the Lowicryl K4M-embedded tissue, the ultrastructure of the neuropil, and particularly that of synapses, was poorly maintained. In contrast, the osmicated epoxy-embedded material offered optimal morphological preservation together with accurate subcellular localization of all antigens under study. The latter approach thus enabled clear visualization of CGRP, tachykinin, and somatostatin immunoreactivity restricted to large dense-cored vesicles (90-150 nm diameter) in many axonal and synaptic profiles in the superficial layers of the dorsal horn. CGRP- and tachykinin-positive profiles were also present in the tract of Lissauer. GABA immunoreactivity was present mainly in axons and terminals, and less frequently in somatic and dendritic profiles. In terminals, which often formed symmetrical synapses on immunonegative dendritic profiles, it was associated with small (30-60 nm diameter) clear vesicles and mitochondria. Double immunolabeling was possible on all preparations, but the osmicated, epoxy-embedded material clearly showed co-localization of peptides, especially of CGRP and tachykinins, within the same dense-cored vesicles in axonal fibers and/or terminals. On the other hand, peptide and GABA immunoreactivity were consistently seen in different nerve profiles. In a few cases, GABAnergic terminals were seen to synapse on tachykinin-positive fibers.

Region-specific appearance of myelin constituents in the developing rat spinal cord

The appearance of myelin-specific glycolipids and of myelin basic protein was studied with regard to the detailed anatomy of the rat cervical spinal cord. The expression of these constituents in particular fibre tracts and regions occurs at specific times of development between postnatal days 1 and 14. This mosaic-like appearance started in the ventral funiculus (day 1) followed by fasciculus cuneatus and ventro-lateral funiculus (day 2), and fasciculus gracilis and dorso-lateral funiculus (days 3 to 4). Cortico-spinal tract (day 11), Lissauer tract (day 14) and the commissures started to acquire myelin very late. In the grey matter, myelin constituents appeared around days 11 to 14 in a patchy pattern. These results support a concept of highly local interactions regulating oligodendrocyte differentiation. In addition, a general rostro-caudal gradient of myelin development exists in the spinal cord, which is independent of the ascending or descending nature of the fibre tracts. Appearance of myelin constitutents in the caudal spinal cord was not prevented by a neonatal transection at mid-thoracic levels.

The plant lectin soybean agglutinin binds to the soma, axon and central terminals of a subpopulation of small-diameter primary sensory neurons in the rat and cat

The distribution of binding of the plant lectin soybean agglutinin to primary sensory neurons has been investigated in the rat and cat. Soybean agglutinin was found to bind to approximately 30% of neurons in the fourth, lumbar dorsal root ganglion of the rat and approximately 50% of neurons in the first, sacral dorsal root ganglion in the cat. Morphometric analysis of these dorsal root ganglia revealed that in both species those neurons which bind soybean agglutinin appear to be a subpopulation of the small-diameter cells. Electron microscopic analysis of the lumbar dorsal roots revealed that soybean agglutinin binds to the plasma membrane of a subpopulation of unmyelinated axons in both species. Myelinated axons did not bind the lectin. In the dorsal horn of the spinal cord, soybean agglutinin bound to components of Lissauer's tract and the superficial laminae (laminae I and II) of the dorsal horn in both species. In the dorsal horn lectin binding was limited to lamina I and the outer portion of lamina II (sublamina II o in the cat, while in the rat lamina I and the entire dorsoventral extent of lamina II (sublaminae II oand II i were labelled. Thirty days following dorsal rhizotomy, soybean agglutinin binding in the superficial dorsal horn, ipsilateral to the rhizotomy, was abolished in both species. These results show the plant lectin soybean agglutinin to be a histochemical marker for the soma and central terminals of a subpopulation of small-diameter sensory neurons in both the rat and cat. It is suggested that soybean agglutinin binding may be used in conjunction with immunohistochemstry to allow identification of putative neurotransmitters within, or in synaptic profiles associated with, the central terminals of small-diameter primary sensory neurons.

Ulex europaeus Agglutinin-I Binding to Dental Primary Afferent Projections in the Spinal Trigeminal Complex Combined with Double Immunolabeling of Substance P and GABA Elements Using Peroxidase and Colloidal Gold

Ulex europaeus agglutinin I (UEA-I) is a plant lectin with an affinity for L-fucosyl residues in the chains of lactoseries oligosaccharides associated with medium- and smaller-diameter dorsal root ganglion neurons and their axonal processes. These enter Lissauer's tract and terminate within the superficial laminae of the spinal cord overlapping projections known to have a nociceptive function. This implies that the surface coatings of neuronal membranes may have a relationship with functional modalities. The present investigation further examined this concept by studying a neuronal projection with a nociceptive function to determine whether fucosyl-lactoseries residues were incorporated in its primary afferent terminals. Transganglionic transport of horseradish peroxidase (HRP) following injection into tooth pulp chambers was employed to demonstrate dental pulp terminals in the trigeminal spinal complex, while peroxidase and fluorescent tags were used concomitantly to stain for UEA-I. Double immunolabeling for substance P (SP) and gamma-aminobutyric acid (GABA) using peroxidase and colloidal gold allowed a comparison of the distribution of a known excitatory nociceptive transmitter with that of UEA-I binding in specific subnuclei. Synaptic interrelationships between UEA-I positive dental pulp primary afferent inputs and specific inhibitory terminals were also examined. SP immunoreactivity occurred in laminae I and outer lamina II (IIo) of subnucleus caudalis (Vc) and in the ventrolateral and lateral marginal region of the caudal half of subnucleus interpolaris (Vi), including the periobex area in which Vi is slightly overlapped on its lateral aspect by cellular elements of Vc. The adjacent interstitial nucleus (IN) also showed an intense immunoreactivity for this peptide antibody. UEA-I binding displayed a similar distribution pattern in both Vc and Vi, but extended into lamina IIi and the superficial part of Lamina III in Vc. Dental pulp terminals were found to have a comparable distribution; however, many extended into the dorsal portion of the caudal half of Vi and the ventromedial quadrant of rostral Vi. Electron-microscopic analysis showed that transganglionically labeled dental pulp terminals contained ovoid, complex membrane-bound vacuoles laden with transported HRP. The preterminal axon and synaptic membranes of those dental pulp terminals located in zones of Vc and Vi displaying an affinity for UEA-I were usually characterized by a patchy, electron-dense coating of the peroxidase tag. SP was demonstrated ultrastructurally with Protein-A colloidal gold (3-nm particles), whereas GABA immunoreactivity was revealed by the avidin-biotin-peroxidase method.(ABSTRACT TRUNCATED AT 400 WORDS)

Distribution of substance P and vasoactive intestinal polypeptide neurons in the chicken spinal cord, with notes on their postnatal development

The distribution of substance P (SP) and vasoactive intestinal polypeptide (VIP) was investigated by immunohistochemistry in the adult chicken spinal cord. By using colchicine treatment, populations of neurons containing either SP or VIP was observed in several regions of the spinal cord. SP neurons were found dorsal to the central canal (CC) and in lamina IV throughout the cord. However, at the thoracic level, numerous relatively larger SP perikarya were located ventral to the CC and aligned on either side of the midline. The distribution of SP fibers is very similar to that reported previously in mammals: they were mostly observed in laminae I and II, in Lissauer's tract, in the dorsolateral funiculus, and dorsal to the CC. In addition, two dense plexuses of SP fibers were noticed in lamina IV. VIP neurons were located mainly in lamina I, in the nucleus of the dorsolateral funiculus, and in the lateral portion of the neck of the dorsal horn throughout the spinal cord. At the thoracic level, many also were located lateral to the CC. Occasionally, single VIP neurons also were encountered dorsal to the CC, in laminae II-IV, and in the intermediate zone. VIP fibers were observed in similar numbers at all spinal levels, occurring mainly in laminae II (probably I) and III, dorsal to the CC, and in the intermediate zone. In addition, examination of the developing chick spinal cords showed similar results as in adult chickens.

The projection of the medial and posterior articular nerves of the cat's knee to the spinal cord

We studied the spinal projections of the medial and posterior articular nerves (MAN and PAN) of the knee joint in the cat with the aid of the transganglionic transport of horseradish peroxidase. The afferent fibers of the MAN entered the spinal cord via the lumbar dorsal roots L5 and L6 and those of the PAN entered via the dorsal roots L6 and L7. Within the dorsal root ganglia, most labeled neurons had small to medium diameters. A relatively higher number of medium-size cell bodies were labeled from the PAN than from the MAN. In the spinal cord labeled MAN afferent fibers and terminations were most dense in the L5 and L6 segments, and those of the PAN were most dense in L6 and L7, that is, in the respective segments of entry. Labeled afferent fibers from both nerves projected rostrally at least as far as L1 and caudally as far as S2. Labeled fibers were found in Lissauer's tract as well as in the dorsal column immediately adjacent to the dorsal horn. In the spinal gray matter, both nerves had two main projection fields, one in the cap of the dorsal horn in lamina I, the other in the deep dorsal horn in laminae V-VI and the dorsal part of lamina VII. Both nerves, but particularly the PAN, projected to the medial portion of Clarke's column. No projection was found to laminae II, III, and IV of the dorsal horn or to the ventral horn. Since these findings parallel observations on hindlimb muscle afferent fibers, the present data support the existence of a common pattern for the central distribution of deep somatic afferent fibers.

Development and distribution of enkephalin-immunoreactive elements in the chicken spinal cord

The development and distribution of methionine-enkephalin-immunoreactive elements were studied in the chicken spinal cord with the indirect immunofluorescence method. Methionine-enkephalin-like immunoreactivity was first detected in the chick spinal cord at embryonic stages 29–30 (incubation day 6). Before stage 35 (day 9). it was mainly observed in fibres almost throughout the while matter. Subsequently, fibres containing the peptide appeared in the ventral half of the gray matter, but mostly in the lateral portion of the neck of the dorsal horn. From stage 40 (day 13 or 14), fibres were especially noticed in laminae 1 and 2. and in the area dorsal to the central canal. In particular, many enkephalin-immunoreactive perikarya were observed in several spinal areas during this period. Such a distribution of both enkephalin-immunoreactive fibres and perikarya remained visible at later embryonic stages, but labelled cells gradually decreased in number and disappeared after hatching. With colchicine treatment, however, a similar distribution of the peptide was found in the spinal cord of adult chickens. As in the embryo, enkephalin-immunoreactive perikarya were mainly observed in the lateral portion of the neck of the dorsal horn, in lamina 1, and in the nucleus of the dorsolateral funiculus throughout the spinal eord. At the thoracic level, many were also located ventral to the central canal. Enkephalin-immunoreactive fibres increased notably in the gray matter of adult chickens. They mainly occurred in laminae 1 and 2, in the lateral portion of the neck of the dorsal horn, and in the area around, especially dorsal to. the central canal. In contrast, enkephalin-immunoreactive fibres decreased in the white matter and they were mainly observed in the dorsolateral funiculus, in Lissauer's tract, and in the lateral funiculus adjacent to the gray. The distribution of enkephalin-immunoreactive fibres was generally comparable at all spinal levels examined. In addition, examination of post-hatched chickens showed virtually the same results as in the adult.

Intraspinal sprouting after administration of nerve growth factor antibodies to neonatal rats

Previous observations show that neonatal removal of nerve growth factor results in an increased number of fine dorsal root axons in the adult animal. The present study shows that the same treatment results in 49% more axons in the adult tract of Lissauer, which is an area of spinal white matter that contains predominant fine primary afferentaxons. These data are consistent with the idea that postnatal removal of NGF leads to increased numbers of fine primary afferent axons which then pass into the white matter of the spinal cord and there make synaptic contacts. If so, this would be a useful preparation for studies on the mechanisms by which newly formed sensory axons can be induced to enter the spinal cord postnatally.

Soybean agglutinin binds to a subpopulation of primary sensory neurones in the cat

The binding of the plant lectin soybean agglutinin (SBA) to primary sensory neurones has been investigated in the rat. SBA binding was found in Lissauer's tract and in laminae I and IIo of the dorsal horn at cervical, thoracic and lumbar levels. Morphometric analysis of the S1 dorsal root ganglia revealed that SBA binding was associated with the small diameter cell population, considered to be the cell bodies of unmyelinated afferent fibres (C-fibres). These findings suggest that SBA may be a useful ultrastructural marker for C-fibre terminals.

Calcitonin gene-related peptide (CGRP) in the human spinal cord: a light and electron microscopic analysis.

The distribution of CGRP immunoreactivity in the cervical, thoracic, lumbar, and sacral levels of the human spinal cord was mapped at the light microscopic level with the aid of a rabbit-generated antiserum against human calcitonin gene-related peptide (CGRP). CGRP-positive fibers formed a dense plexus in lamina I, II, the reticulated region of lamina V, and the tract of Lissauer at all spinal cord levels. The distribution of fibers showed some variations dependent on the cord level analyzed. At the light microscopic level, intervaricose fiber diameters consistently measured 1.0 micron or less, and two types of CGRP varicosities were observed: a small (1 to 2 microns in diameter), relatively round profile and a larger, (3 to 4 microns in diameter) oval or oblong profile. At the electron microscopic level, immunostained varicosities contained a mixture of round clear vesicles and vesicles that contained dense cores. The CGRP immunoreaction product was often associated with vesicles containing dense cores. The reaction product was also seen associated with clear round vesicles or in the cytoplasmic matrix. Postsynaptic elements included dendritic spines, small and large diameter dendritic shafts and vesicle containing profiles. The presence of CGRP in the superficial dorsal horn of human spinal cord is highly suggestive of a role in primary afferent transmission as postulated in lower vertebrates. This study establishes the distribution of CGRP at four different spinal levels in human cord and will serve as a basis for future studies related to the pathologic conditions affecting sensory systems.

A light and electron microscopic study of calcitonin gener-related peptide in the spinal cord of the rat

The present study localized calcitonin gene-related peptide at the light and electron microscopic levels in the lumbar spinal cord of the rat. One finding was that axons and terminals were labeled in both lamina I and Ilo medially but only in lamina I laterally. The functional implications of this innervation pattern are not clear but presumably this anatomic arrangement bears on both dorsoventral and mediolateral patterns of organization of primary afferent input into the dorsal horn. We also found that although the means of labeled myelinated and unmyelinated axon diameters in the tract of Lissauer were different, there was great overlap in these populations. Furthermore, subcellular localizations indicated that immunostaining of calcitonin gene-related peptide was associated primarily with microtubules in axons and cores of large dense-core vesicles in presynaptic terminals. Finally, labeled presynaptic terminals contained relatively few large dense-core vesicles and formed the presynaptic elements of simple axodendritic contacts almost exclusively. These last findings contrast with localizations of calcitonin gene-related peptide in the monkey, which has many more large dense-core vesicles in labeled terminals and in which a much higher proportion of labeled endings form the central parts of glomeruli.

The spinal projection of individual identified A-delta- and C-fibers.

1. Recordings were made from individual sensory neurons with an A-delta peripheral conduction velocity, either intrasomally in the L7 dorsal root ganglion, or extracellularly in Lissauer's Tract or in the dorsal root close to the root entry zone. The spinal projection of these afferents was assessed by their antidromic response to stimulation of the dorsal columns (DC) or Lissauer's Tract (LT) at the L5/L6 border. The adequate stimulus was also ascertained. 2. A-delta-fibers could be divided into two groups: high-threshold mechanoreceptors from either skin or muscle (HTMRs) and low-threshold mechanoreceptors (LTMs), primarily Down Hairs. A third group of cells recorded intrasomally had broad spikes with shoulders on the downstroke characteristic of A-delta-nociceptors and were so classified provisionally, although no adequate stimulus could be identified. HTMRs and broad spike cells projected either in DC or LT, but LTMs projected only in DC, never in LT. About one-quarter of both groups failed to project rostrally as far as L5/L6. 3. Cells with unmyelinated axons recorded intrasomally were found to supply either low-threshold or high-threshold mechanoreceptors. Unlike A-delta-cells, all these cells had broad spikes with shoulders on the downstroke. Proportionally fewer C-fibers than A-delta-fibers projected as far as one segment rostral from their root entry zone. Of those that did, axons supplying low-threshold mechanoreceptors projected only in DC, whereas those innervating high-threshold mechanoreceptors could project either through LT or DC. 4. A-delta-fibers supplying LTMs and HTMRs exhibited a similar reduced conduction velocity was reduced even further in the spinal cord but much more for HTMRs than for LTMs. For C-fibers the conduction velocity decrease was more substantial in the dorsal root for HTMRs than for LTMs. 5. These findings suggest that axons innervating different peripheral receptors exhibit characteristic cellular properties. They confirm that the primary afferent component of Lissauer's Tract is specialized as a "pain pathway" but also indicate that the dorsal columns may play some role in the transmission of nociceptive information.

Pain control with laser-produced dorsal root entry zone lesions.

Pain relief was evaluated in 40 patients with various types of deafferentation pain that were treated with dorsal root entry zone (DREZ) lesions produced with microsurgical lasers. Good long-term pain relief was evident in some paraplegics and in all patients with brachial plexus avulsion. Several other small subgroups of patients benefited from laser DREZ lesions as well. Pain associated with arachnoiditis and peripheral nerve injury or neuropathy did not respond to laser DREZ lesioning. Based upon the smaller lesion dimensions produced with the lasers, it is proposed that interruption of impulses in the tract of Lissauer may be a mechanism of pain control in patients that responded to laser DREZ lesions.

Normal anatomy and physiology of the spinal cord dorsal horn.

The dorsal horn of the spinal cord receives afferent input from innocuous primary afferent neurons via collaterals from the dorsal columns. This input is integrated and relayed primarily by neurons in laminae III-VI. Dorsal horn neurons which encode innocuous inputs project to the medulla and the cervical spinal cord via the dorsal columns and the dorsolateral funiculus. Nociceptive primary afferent neurons enter the spinal dorsal horn via collaterals from Lissauer's tract. Nociceptive input is integrated and relayed by neurons in laminae I, II and V which project to the reticular formation and thalamus via the anterolateral tract.

Organization of glutamate‐like immunoreactivity in the rat superficial dorsal horn: Light and electron microscopic observations

Glutamate has been shown to be a neurotransmitter in the central nervous system of vertebrates, and it has been hypothesized that glutamate is functional as a neurotransmitter in the spinal cord dorsal horn. A monoclonal antibody to fixative-modified glutamate was used in this study to examine the light microscopic and ultrastructural profiles of glutamate-like immunoreactivity in the superficial dorsal horn of the rat spinal cord. Glutamate-like immunoreactivity was observed in neurons, fibers, and terminals of both laminae I and II. Marginal zone immunoreactive neurons ranged from 10 to 30 micron in diameter and received many nonimmunoreactive somatic synapses. In substantia gelatinosa, immunoreactive neurons were observed in both inner and outer layers, ranged 5 to 10 micron in diameter, and received few nonimmunoreactive somatic synapses. Glutamate-like immunoreactive dendrites were observed in both laminae and were contacted primarily by nonimmunoreactive synaptic terminals that generally contained small clear vesicles. Both myelinated and unmyelinated immunoreactive axons were observed in Lissauer's tract. Immunoreactive terminals contained small (40 nm) clear vesicles and generally formed simple synaptic contacts with nonimmunoreactive dendrites in laminae I and II. The results of this study corroborate the importance of glutamate as a neurotransmitter in spinal sensory mechanisms.

Substance P-like immunoreactive fibers in the trigeminal sensory nuclei of the pit viper, Trimeresurus flavoviridis

Substance P-like immunoreactive nerve fibres were located in the trigeminal sensory system of the infrared-sensitive snake, Trimeresurus flavoviridis, using the immunohistochemical method. There are two trigeminal sensory systems in the medulla of this animal: the descending nucleus and the lateral descending nucleus. The descending nucleus is equivalent to the trigeminal spinal nucleus in other vertebrates, and the lateral descending nucleus is a special trigeminal sensory nucleus belonging to the infrared sensory system. In the present study we determined that the lateral descending nucleus is completely ensheathed by large numbers of substance P-like immunoreactive fibers. The distribution of these fibers seems to be similar to that of the thin vagal unmyelinated fibers, rather than to that of the thick trigeminal myelinated fibers. More substance P-like immunoreactive nerve fibers were observed in the lateral descending tract than in the descending tract. Almost no dense substance P-like immunoreactive fibers were found in these tracts rostral to the lateral descending nucleus or rostral to the subnucleus caudalis of the descending nucleus. The substance P-like immunoreactive fibers in the lateral descending tract extended to those of Lissauer's tract of the spinal cord, and the substance P-like immunoreactive fibers surrounding the Lissauer's tract were similar in appearance to those of the lateral descending nucleus. This nucleus seems to have developed from the elements existing in Lissauer's tract, and also to have a similar modulating function. The primary nucleus of the infrared sensory system is the most substance P-like immunoreactive nucleus in the trigeminal sensory system of this animal. Even in the trigeminal sensory system, substance P-like immunoreactive fibers seem not to be related solely to the nociceptive sensation.