No abstract

Two major interneurons of the outer part of Rexed's layer I1 (IIa) were impaled with microelectrodes, had their primary inputs characterized, and were subsequently filled with horseradish peroxidase. Their fine structural characteristics and synaptic connections were then analyzed electron microscopically. Two islet cells, whose rostrocaudally oriented dendrites were largely confined within layer IIa, received primary input from small myelinated axons. A stalked cell, whose cell body was situated on 1/11 border had a cone-shaped dendritic arbor which traversed layer IIa as well as the inner part of Rexed's layer I1 (IIb) and rostrocaudal dendritic branches which ran for part of their course along the UIIa border. It received primary input from small myelinated as well as from unmyelinated axons. Both cell types received asymmetrical axodendritic synapses from primary endings in layer IIa and IIb glomeruli and widely separated symmetrical axodendritic synapses from small nonprimary endings outside of glomerli.The presence of aggregates of synaptic vesicles in the dendrites of the layer IIa islet cells but not in the stalked cell dendrites constitutes the major fine structural difference between these interneurons. Islet cell dendrites form symmetrical synapses on several different kinds of neural processes. They usually send either a single type 2 spine (spines which contain synaptic vesicles) or dendritic shaft into layer IIa and IIb glomeruli, where they form dendrodendritic synapses on adjacent type 1 spines (spines without synaptic vesicles) and on other small dendritic shafts. Some islet cell type 2 spines also form dendroaxonic synapses on the primary endings.Outside of the glomeruli, islet cell dendrites also form dendrodendritic synapses on type 1 spines and different sized dendritic shafts. They often approach other dendritic shafts forming small bundles of dendrites in which they are reciprocally linked by dendrodendritic synapses to other synaptic vesicle-containing dendrites. At bead-like enlargements of their dendritic shafts and along some of their fine caliber dendritic shafts, the islet cells form dendroaxonic synapses on the shafts of unmyelinated axons. The unmyelinated axon of the islet cell forms symmetrical synapses on layer I1 dendritic shafts and spines outside of glomeruli. The role of the layer IIa islet cell as an inhibitory interneuron is discussed.

Neurons in Rexed's lamina I have the bulk of their dendritic arbors confined within this lamina. This study examines the morphology and synaptic connections of primary axons which generate axonal endings in lamina I of the spinal dorsal horn and are in position to deliver their inputs directly to lamina I neurons. Primary axons were made visible for light and electron microscopical study by applying horseradish peroxidase (HRP) to the severed central stumps of cervical and lumbar dorsal roots and allowing sufficient time for the orthograde movement of the HRP into the terminal axonal arbors. Golgi preparations provided supplementary light microscopical views of these axons. Lamina

Recent studies have suggested that calcitonin gene-related peptide (CGRP) can be used as a marker for a subpopulation of nociceptive primary afferents. Consequently, CGRP-immunoreactive (CGRP-IR) primary afferents have been reported to project many segments rostral to their segment of entry and to send collaterals into the superficial and deep laminae of the dorsal horn. This study reports that some CGRP-IR primary afferents of sacral origin project rostral through the ipsilateral lumbar enlargement in the cat. The ultrastructure of these multisegmentally projecting primary afferent axons and terminals identified in a partially denervated cat was examined and compared to the ultrastructure of CGRP-IR afferents from an intact cat. Retrograde transport of wheatgerm agglutinin-colloidal gold injected into the cat L4 spinal cord resulted in labeling of primary afferent cell bodies in the ipsilateral L4-S1 dorsal root ganglia (DRG). Analysis of every fourth section through the ipsilateral S1 DRG revealed as many as 1,000 retrogradely labeled neuronal cell bodies. One third of these cell bodies were double labeled for CGRP-like immunoreactivity. The number of single- and double-labeled cells increased in ganglia closer to the injection site (L4-L7). At the ultrastructural level, in the lumbosacral dorsal spinal cord of a normal cat, most CGRP-IR axons were unmyelinated, while the rest were small myelinated axons. In both the superficial dorsal horn and lamina V, CGRP-IR varicosities were dome shaped, scallop shaped, or elongated. The CGRP-IR varicosities contained small agranular vesicles and frequently a few dense core vesicles. These labeled varicosities formed asymmetric synapses on unlabeled dendritic spines, shafts, or neuronal somata. One cat received multiple unilateral dorsal rhizotomies (S1-L4) and an ipsilateral hemisection (mid L4). CGRP-IR axons and terminals were found within each of the rhizotomized segments, although their density was greatly reduced compared to that in the intact animals. In Lissauer's tract the majority (greater than 90%) of CGRP-IR fibers were unmyelinated. In laminae I and V, the remaining CGRP-IR varicosities were mainly the dome-shaped varicosities morphologically similar to those observed in the normal spinal cords. They contained small agranular vesicles and a few dense core vesicles and formed asymmetric synapses on unlabeled dendritic shafts and spines. These data demonstrate that unmyelinated, presumably C-fiber nociceptive primary afferents and some small myelinated A-delta nociceptive primary afferents of sacral origin project rostral through the cat lumbar enlargement and make synaptic connections in both the superficial and deep laminae of the cat dorsal spinal cord.(ABSTRACT TRUNCATED AT 250 WORDS)