William M. Falls

Periventricular-Hypophysial Dopaminergic Neurons Innervate the Intermediate but Not the Neural Lobe of the Rat Pituitary Gland

The purpose of the present study was to determine the relative distribution of axon terminals of A14 periventricular-hypophysial dopaminergic (PHDA) neurons in the neural and intermediate lobes of the rat pituitary gland. Discrete unilateral injections of the anterograde tracer Phaseolus vulgaris leucoagglutinin (PHA-L) into the periventricular nucleus resulted in labelling of extensively branched terminal axonal arbors in the intermediate lobe, but not the neural lobe of the pituitary gland. In contrast, unilateral injections of PHA-L into the paraventricular nucleus revealed thick, varicose terminal arborizations containing PHA-L in the neural lobe, but not the intermediate lobe. Terminal axonal branches and varicosities containing PHA-L immunoreactivity in the intermediate lobe were also immunoreactive for tyrosine hydroxylase. These results reveal that A14 PHDA neurons originating in the periventricular nucleus of the hypothalamus project axons to the intermediate lobe of the rat pituitary gland.

The Interstitial System of the Spinal Trigeminal Tract in the Rat: Anatomical Evidence for Morphological and Functional Heterogeneity

Utilizing cyto-, myelo-, and chemoarchitecture as well as connectional criteria, the present study reveals the interstitial system of the spinal trigeminal tract (InSy-SVT) in the rat to be composed of five morphologically and functionally distinct components that are distributed within spatially restricted regions of the lateral medulla. The first component is represented by scattered interstitial cells and neuropil, which extend laterally into SVT from the superficial laminae of the medullary dorsal horn (MDH). The second component, the dorsal paramarginal nucleus (PaMd), consists of a small group of marginal (lamina I)-like neurons and neuropil situated within the dorsolateral part of SVT at the rostral pole of MDH. The third component represents a trigeminal extension of the parvocellular reticular formation (V-Rpc) into the ventromedial aspect of SVT at levels extending from rostral MDH to the caudal part of trigeminal nucleus interpolaris (Vi). The fourth component, the paratrigeminal nucleus (PaV), consists of a large accumulation of neurons and neuropil situated within the dorsal part of SVT throughout the caudal half of Vi. The fifth component is the insular trigeminal-cuneatus lateralis nucleus (iV-Cul), which is a discontinuous collection of neurons and neuropil interspersed among fibers of SVT as well as wedged between it and the spinocerebellar tract. Thalamic projection neurons are located in PaMd and V-Rpc, whereas cerebellar projecting neurons are confined to iV-Cul.

The dorsomedial portion of trigeminal nucleus oralis (Vo) in the rat: cytology and projections to the cerebellum.

Electrophysiological studies have described four major tactile areas in the rat cerebellar cortex. These areas are in crus I, crus II, the paramedian lobule (PML), and the uvula, and a major portion of each is related to the ipsilateral orofacial region. This study demonstrates that neurons in trigeminal nucleus oralis (Vo) that project to the orofacial portions of these four major tactile areas are localized in the dorsomedial (DM) subdivision of the nucleus. The distribution, light-microscopic morphology, and relative densities of trigeminocerebellar neurons within DM, retrogradely labeled with horseradish peroxidase (HRP) following injections into each of the four major tactile areas, were analyzed and compared as well as correlated with the myelo- and cytoarchitecture of DM observed in Nissl sections, 1-micron sections, and Golgi material. On the basis of myelo- and cytoarchitectonic as well as trigeminocerebellar connectional criteria, three portions of DM were identified: caudal DM (CDM), middle DM (MDM), and rostral DM (RDM). The greatest portion of DM is made up of MDM (1.3 mm long), which can be further subdivided into dorsal (MDMd) and ventral (MDMv) zones. CDM forms the caudal 800 microns of DM, while RDM makes up the rostral 280 microns of the subdivision. Longitudinally running deep axon bundles permeate CDM, MDMv, and RDM, but are conspicuously absent from MDMd. The majority of neurons found throughout CDM, MDMv, and RDM have medium-sized (15- to 30-microns) somata and can be divided into two types on the basis of their somatodendritic morphology. CDM, MDMv, and RDM also contain a small neuronal cell type (5- to 15-microns cell body) that is encountered less frequently than either one of the two types of medium-sized cells. A fourth type of neuron with a large (25- to 50-microns) fusiform- to pyramidal-shaped cell body is the least frequently observed neuronal cell type and is located principally in CDM and MDMv. MDMd contains a fifth type of neuron characterized by a small (5- to 15-microns) oval soma. Data from the retrograde HRP experiments show that all five of these neuronal cell types in their respective portions of DM project to one or more of the orofacial portions of the four major tactile areas of the cerebellar cortex. Many medium-sized neurons of both types in CDM, MDMv, and RDM project to crus I, crus II, and/or PML.(ABSTRACT TRUNCATED AT 250 WORDS)

The morphology of neurons in trigeminal nucleus oralis projecting to the medullary dorsal horn (trigeminal nucleus caudalis): A retrograde horseradish peroxidase and golgi study

This study demonstrates that trigeminal nucleus oralis, the most rostral subdivision of the spinal trigeminal nucleus, contains four morphologically distinct types of small neurons which project to the medullary dorsal horn (trigeminal nucleus caudalis) via descending intratrigeminal pathways. Using the retrograde transport of horseradish peroxidase following injections in the medullary dorsal horn, labeled small neurons with cell bodies ranging from 8-15 microns in diameter are found principally in the ventrolateral portion of the trigeminal nucleus oralis. Most neurons are labeled ipsilaterally throughout the entire rostrocaudal extent of the ventrolateral portion of the trigeminal nucleus oralis, but a few cells are also labeled contralaterally. From this aspect of the present study it can be concluded that a specific portion of the trigeminal nucleus oralis, i.e. the ventrolateral part, contains numerous small neurons which send descending projections to the medullary dorsal horn that could affect synaptic activity there. Utilizing both the methods of Golgi and retrograde horseradish peroxidase labeling four distinct types of small descending medullary dorsal horn projection neurons can be distinguished in the ventrolateral portion of the trigeminal nucleus oralis on the basis of their morphology and the distribution of their axons and dendrites. All four neuronal cell types are present throughout the entire rostrocaudal extent of the trigeminal nucleus oralis. Type I neurons are the most frequently labeled descending medullary dorsal horn projection neurons. They are concentrated in the medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis and display dendritic trees which occupy spherical domains approaching 300 microns in diameter. The unmyelinated axons of many of these cells arise either directly from the cell body or a primary dendrite and give rise to a single collateral within 50 microns of their site of origin. This collateral generates a fine axonal plexus within a portion of the dendritic arbor of the parent cell while the parent axon, without branching further, travels a short distance in the ventrolateral portion of the trigeminal nucleus oralis and enters a deep axon bundle. Type II neurons are the second most frequently labeled descending medullary dorsal horn projection neuron. They generate medial and lateral dendritic arbors which together span nearly the entire medial 500-550 microns of the ventrolateral portion of the trigeminal nucleus oralis. An unmyelinated axon emerges from the cell body and within 10-30 microns of its origin gives rise to two collaterals.(ABSTRACT TRUNCATED AT 400 WORDS)

Termination in trigeminal nucleus oralis of ascending intratrigeminal axons originating from neurons in the medullary dorsal horn: an HRP study in the rat employing light and electron microscopy

The anterograde horseradish peroxidase (HRP) technique was used to identify ascending intratrigeminal axons originating from neurons in the medullary dorsal horn (MDH) which terminate in trigeminal nucleus oralis (Vo). HRP injections into the MDH labeled two populations of axons ascending ipsilaterally within the spinal trigeminal nucleus. The first population was composed of parent branches which each gave off a single branching collateral strand to Vo as they ascended. These collaterals were characterized by boutons filled with small, round synaptic vesicles and forming asymmetrical synaptic contacts with large diameter dendritic shafts. The second axonal population was made up of parent branches which terminated directly in Vo. Their short terminal strands were distinguished by axonal endings containing pleomorphic synaptic vesicles and forming symmetrical synaptic junctions with small diameter dendritic shafts and spines.

The synaptic organization of the cerebello-oiivary circuit

SummarySection of the superior cerebellar peduncle just rostral to the deep cerebellar nuclei results in degenerating axon terminals within the contralateral inferior olive. The nuclear origin of this fiber system and its distribution within the subdivisions of the inferior olive were described in a companion study (Martin et al., 1976). Precise localization of these degenerating terminals within the nucleus was accomplished by the examination of 1 μ plastic sections cut from each tissue block prior to thin sectioning. Degenerating axon terminals are present in all the nuclear subdivisions and when seen with the electron microscope they frequently are localized in the previously described synaptic clusters (King, 1976). These terminals demonstrate an electron dense reaction at survival times of 2 and 3 days. By day 4, they are shrunken and irregular in shape, and typically are surrounded by astrocyte processes. Cerebello-olivary axon terminals measure 1–3 μ, contain spherical, clear synaptic vesicles and typically contact spiny appendages within the synaptic clusters (glomeruli). Thus, we have demonstrated that one of the primary axon systems which terminates within the synaptic clusters is from the cerebellar nuclei. We have yet to determine the origins of the remaining terminals within the synaptic clusters which include endings with either smaller spherical, pleomorphic or numerous dense core vesicles.

A comparison of the distribution and morphology of thalamic, cerebellar and spinal projection neurons in rat trigeminal nucleus interpolaris.

The retrograde transport of horseradish peroxidase was used to examine and compare the distribution and morphology of thalamic, cerebellar and spinal projecting neurons in rat trigeminal nucleus interpolaris following large injections into their respective targets. The regional distribution of these three populations was evaluated in relation to the six cytoarchitecturally distinct regions which characterize the nucleus. Cerebellar projecting neurons were distributed throughout the rostrocaudal extent of trigeminal nucleus interpolaris, but were infrequently present in its dorsolateral region and in the rostral pole of the nucleus. Thalamic projecting neurons exhibited a distribution pattern that extensively overlapped with that of the trigeminocerebellar neurons: however, they were particularly concentrated in caudal, dorsomedial and rostral, ventrolateral regions of the nucleus. Trigeminospinal projecting neurons exhibited a more restricted distribution within ventral and lateral regions of trigeminal nucleus interpolaris. Although the three populations of projection neurons could not be distinguished solely on the basis of somatic size or shape, distinct regional variations in the distribution and somatodendritic and axonal morphology of these neurons indicated that they arise largely from independent cell populations. However, several regions were identified in which specific cell types were likely to contribute to axonal collaterilization among these pathways. In the ventrolateral magnocellular region of the nucleus, for example, more than half of the large multipolar-shaped neurons were retrogradely labeled after injections into each of the three target sites. The results of the present study indicate that the thalamic, cerebellar and spinal projections of trigeminal nucleus interpolaris arise from a morphologically heterogeneous group of neurons. In addition, regional variations in the distribution and morphology of these neurons provide evidence for the existence of functionally distinct regions that parallel the cytoarchitecturally defined regions of the nucleus. This study also provides indirect evidence for and against collateralization among these three projections within specific regions of the nucleus.

An AnAlysis of the Cyto- and Myeloarchitectonic Organization of Trigeminal Nucleus Interpolaris in the Rat

The cyto- and myeloarchitectonic organization of trigeminal nucleus interpolaris (Vi) was examined in the rat using correlated Nissl- and myelin-stained sections. The caudal boundary of Vi is marked by a spatial overlap with the rostral pole of the medullary dorsal horn (MDH), where there is a dorsal and medial displacement of the substantia gelatinosa (SG, lamina II) layer of MDH. This spatial displacement was further documented using cytochrome-oxidase-reacted sections through the periobex region (POR) of the medulla, where the relatively unstained SG contrasts sharply with the intensely stained Vi neuropil. The rostral boundary of Vi is characterized partly by a distinct overlap with the caudal pole of the dorsomedial region (DM) of trigeminal nucleus oralis (Vo), and partly by a more gradual transition with ventral and lateral regions of Vo. The presence of the distinct MDH-Vi overlap is discussed in terms of its impact on the widespread contention that Vi is involved in the processing of dental pain afferents in the POR. Six separate and distinct regions of rat Vi can be distinguished on the basis of differences in their overall cyto- and myeloarchitecture: (1) a ventrolateral parvocellular region (vlVipc), which occupies the ventrolateral caudal half of Vi; (2) a ventrolateral magnocellular region (vlVimc), which occupies a similar region in the rostral half of the nucleus; (3) a border region (brVi), interposed between the spinal trigeminal tract (SVT) and vlVipc and vlVimc; (4) a dorsolateral region (dlVi), which lies predominantly in the rostral two-thirds of Vi subjacent to the dorsal half of SVT; (5) a dorsal cap region (dcVi), occupying the dorsomedial aspect of the nucleus throughout its entire rostrocaudal extent; and (6) an intermediate region (irVi), which lies immediately ventral to dcVi within the concavity formed by the medial borders of vlVipc and vlVimc. It is proposed that these cyto- and myeloarchitecturally distinct regions of Vi may largely represent functionally distinct regions, based on reported differences in the organization of afferent and efferent projections within the nucleus.

The spinotrigeminal pathway and its spatial relationship to the origin of trigeminospinal projections in the rat

The anterograde transport of horseradish peroxidase and tritiated amino acids was used to examine the distribution and morphology of spinal afferent fibers terminating in the rat spinal trigeminal complex. The results confirm the existence of a direct, ipsilateral projection from the spinal cord which is distributed exclusively to the deepest layers of the medullary dorsal horn narrow regions subjacent to the spinal trigeminal tract in trigeminal nucleus interpolaris, trigeminal nucleus oralis and the trigeminal main sensory nucleus. Spinal inputs also terminated in the insular trigeminal-cuneatus lateralis nucleus which is a distinct component of the interstitial system of the spinal trigeminal tract. The spinal afferent fibers which terminated in the dorsolateral parts of the spinal trigeminal complex arose from the dorsal column funiculi, while those that terminated in ventral parts of the complex arose from both the dorsal column and lateral funiculi. The tritiated amino acid experiments indicate that at least part of the spinotrigeminal pathway originates from cells located in the cervical spinal dorsal horn. The present findings also document a complex spatial relationship between the spinotrigeminal and trigeminospinal pathways which includes an extensive overlap between spinotrigeminal fibers and spinal projecting neurons in each of the lateralmost regions of the complex. This spatial overlap supports the existence of anatomical substrates which may underlie functional reciprocal loops between the spinal trigeminal complex and cervical spinal cord. Since these regions are primarily concerned with the processing of sensory information from lateral and posterior parts of the face, it follows that the spinotrigeminal pathway may be primarily concerned with the integration of head and neck functions. In addition, the spatial convergence of spinal inputs and the distribution of other trigeminal efferent neurons suggests that part of the spinotrigeminal pathway may be involved in spino-trigemino-thalamic and spino-trigemino-cerebellar pathways in parallel with other spinobulbar pathways in the medulla.

Axonal endings terminating on dendrites of identified large trigeminospinal projection neurons in rat trigeminal nucleus oralis

The retrograde horseradish peroxidase technique was used to: (1) identify and assess the overall morphology of large neurons in the ventrolateral portion (VL) of rat trigeminal nucleus oralis projecting to cervical, thoracic and lumbosacral levels of the spinal cord; and (2) characterize the synaptic endings terminating on their dendrites. The morphology of large VL neurons projecting to all spinal levels is similar. They have 25-50 microns pyramidal-shaped somata which emit 3-6 primary dendrites. These primary dendrites give rise to spherical to elliptical-shaped dendritic arbors measuring up to 700 microns in diameter. Labeled axons enter either a deep axon bundle or the medial portion of the spinal V tract. Dendrites of labeled neurons are contacted by axonal endings of 3 types. The most numerous endings are filled with clear, spherical synaptic vesicles and usually form single asymmetrical contacts along the entire length of dendritic shafts. Synapsing less frequently on dendritic shafts are endings containing pleomorphic synaptic vesicles and forming single symmetrical synaptic contacts. The least frequently encountered synaptic terminal contains flattened synaptic vesicles and makes a single symmetrical synaptic contact with a dendritic shaft.