Variation of dorsal horn cell dendritic spread with map scale.

Cells in laminae III, IV, and V of cat dorsal horn were injected with horseradish peroxidase or neurobiotin. Dorsal views of the dendritic domains were constructed in order to measure their lengths, widths, areas, and length/width ratios in the horizontal plane (the plane of the somatotopic map). Dendritic domain width and area in the horizontal plane were negatively correlated with fractional distance between the medial and lateral edges of the dorsal horn. These results are consistent with the hypothesis that dendritic domain width varies with map scale, which is maximal in the medial dorsal horn. This is similar to the variation in widths of primary afferent bouton distributions. The parallel variation of dorsal horn cell dendritic domain width and primary afferent bouton distribution width with map scale suggests that there is a causal relation between morphology and map scale in the dorsal horn representation of the hindlimb. This variation of adult morphology with map scale must reflect mechanisms responsible for the assembly of receptive fields.

Influence of map scale on primary afferent terminal field geometry in cat dorsal horn.

1. Thirty-one physiologically identified primary afferent fibers were labeled intracellularly with horseradish peroxidase (HRP). 2. A computer analysis was used to determine whether the distribution of cutaneous mechanoreceptive afferent terminals varies as a function of location within the dorsal horn somatotopic map. 3. An analysis of the geometry of the projections of these afferents has shown that 1) terminal arbors have a greater mediolateral width within the region of the foot representation than lateral to it, 2) terminal arbors have larger length-to-width ratios outside the foot representation than within it, and 3) the orientation of terminal arbors near the boundary of the foot representation reflects the angle of the boundary. Previous attribution of mediolateral width variations to primary afferent type are probably in error, although there appear to be genuine variations of longitudinal extent as a function of primary afferent type. 4. Nonuniform terminal distributions represent the first of a three-component process underlying assembly of the monosynaptic portions of cell receptive fields (RFs) and the somatotopic map. The other two components consist of the elaboration of cell dendritic trees and the establishment of selective connections. 5. The variation of primary afferent terminal distributions with map location is not an absolute requirement for development of the map; for example, the RFs of postsynaptic cells could be assembled with the use of a uniform terminal distribution for all afferents, everywhere in the map, as long as cell dendrites penetrate the appropriate portions of the presynaptic neuropil and receive connections only from afferent axons contributing to their RFs.(ABSTRACT TRUNCATED AT 250 WORDS)

Somatotopy of digital nerve projections to the cuneate nucleus in the monkey.

Somatotopic arrangements of cells and fibers within the dorsal columns and the dorsal column nuclei have been mapped most precisely by electrophysiological recording methods. This study uses an anatomical approach to evaluate the precision of individual digital nerve projections to the cuneate nucleus (CN) in young macaque monkeys. Digital nerves supplying about one-half the palmar skin of a digit were surgically exposed, cut, and treated with wheatgerm agglutinin conjugated to horseradish peroxidase (WGA:HRP) on 3 successive days. After 2 additional days, animals were killed and medullas were recovered for study of serial sections reacted to display axons labeled by transganglionic transport of label. Labeled afferent fibers from each digit were found within a circumscribed columnar zone extending through the caudal CN and rostrally throughout the pars rotunda of CN. At caudal levels, diffuse projections reach the dorsal edge of the CN; more rostrally, they shift into deeper parts of the nucleus and are heaviest along its ventral and medial edges at levels near the obex. Fibers from the thumb (digit 1) project lateral (and ventral) to those from digit 2, and projections from digit 3 are medial to those from 2. Each digital projection field is closely adjacent to that from the adjacent digit. Few fibers extend to the rostral CN. Projection fields of homologous digits are quite symmetrical on the two sides. Although there do seem to be some differences in the somatotopic arrangement of digital input in macaques compared to other nonprimate mammals studied previously, these observations (precisely organized, circumscribed fields for separate digits) define a system well designed for transmission of data encoding spatial relationships.

Opossum adrenal medulla: I. Postnatal development and normal anatomy.

The anatomy and histology of the adrenal gland in the adult opossum were found to be typical for mammals. The development of the adrenal medulla was also found to follow the typical mammalian pattern. Primitive sympathetic cells were found in both intra- and extra-adrenal locations in the newborn at a time when chromaffin precursor cells were migrating to the adrenal anlage. Pheochromoblasts first appeared within the forming medulla where at a later stage chromaffin cells could be observed forming columns of cells between adjacent sinusoids. Unlike in other mammals, much of this development takes place postnatally when the neonate is in the mother's marsupium. The value of the developing opossum adrenal medulla as an experimental model is stressed, since a significant amount of development takes place in an environment that is accessible to experimental manipulation.

Projection of cervical dorsal root fibers to the medulla oblongata in the brush-tailed possum, Trichosurus vulpecula.

This study describes the projection of cervical spinal afferent nerve fibers to the medulla in the brush-tailed possum, a marsupial mammal. After single dorsal roots (between C2 and T1) were cut in a series of animals, the Fink-Heimer method was used to demonstrate the projection fields of fibers entering the CNS via specific dorsal roots. In the high cervical spinal cord, afferent fibers from each dorsal root form a discrete layer in the dorsal funiculus. The flattened laminae from upper cervical levels are lateral and those from lower cervical levels are medial within the dorsal columns. All afferent fibers at this level are separated from gray matter by the corticospinal fibers in the dorsal funiculus. All cervical roots project throughout most of the length of the well-developed main cuneate nucleus in a loosely segmentotopic fashion. Fibers from rostral roots enter more lateral parts of the nucleus, and fibers from lower levels pass to more medial areas; but terminal projection fields are typically large and overlap extensively. At more rostral medullary levels, fibers from all cervical dorsal roots also reach the external cuneate nucleus. The spatial arrangement here is more complex and more extensively overlapped than in the cuneate nucleus. Rostral cervical root fibers reach ventral and ventrolateral areas of the external cuneate nucleus and continue to its rostral pole; more caudal root fibers project to more dorsal and medial regions within the nucleus. These results demonstrate that projection patterns of spinal afferents in this marsupial are similar to those seen in the few placental species for which detailed data concerning this system are available.

"High spinal" dysrhaphism. Case report of a complex cervical meningocele.

"High spinal" (cervical and upper thoracic) dysrhaphism usually involves either a meningocele or a dermal sinus tract. These high spinal lesions can have a complex intradural anatomy at the level of the lesion (as this case reports) and are associated with an increased incidence of lower spinal occult dysrhaphic anomalies. It is therefore recommended that patients with "high spinal" dysrhaphism undergo radiological evaluation of the entire spine to identify those patients with intradural anomalies, define the anatomy for surgery, and investigate the lower spine for associated occult anomalies.

Efferent projections of the basolateral amygdala in the opossum, Didelphis virginiana.

The autoradiographic anterograde axonal transport technique was used to study efferent projections of the opossum basolateral amygdala. All nuclei of the basolateral amygdala send topographically organized fibers to the bed nucleus of the stria terminalis (BST) via the stria terminalis (ST). Injections into rostrolateral portions of the basal nuclei label fibers that surround the commissural bundle of the ST, cross the midline by passing along the outer aspect of the anterior commissure, and terminate primarily in the contralateral BST, anterior subdivision of the basolateral nucleus (BLa), ventral putamen, and olfactory cortex. Each of the basal nuclei project ipsilaterally to the anterior amygdaloid area, substantia innominata and topographically to the ventral part of the striatum and adjacent olfactory tubercle. The posterior subdivision of the basolateral nucleus (BLp), but not the basomedial nucleus (BM), projects to the ventromedial hypothalamic nucleus. BLa and BLp have projections to the nucleus of the lateral olfactory tract and also send fibers to the central nucleus, as does the lateral nucleus (L). The lateral nucleus also has a strong projection to BM and both nuclei project to the amygdalo-hippocampal area. BLa and BLp send axons to the ventral subiculum and ventral lateral entorhinal area whereas L projects only to the latter area. The lateral nucleus and BLp project to the perirhinal cortex and the posterior agranular insular area. The BLa sends efferents to the anterior agranular insular area. Rostrally this projection is continuous with a projection to the entire frontal cortex located rostral and medial to the orbital sulcus. All of the nuclei of the basolateral amygdala project to areas on the medial wall of the frontal lobe that appear to correspond to the prelimbic and infralimbic areas of other mammals. Despite the great phylogenetic distance separating the opossum from placental mammals, the projections of the opossum basolateral amygdala are very similar to those seen in other mammals. The unique frontal projections of the opossum BLa to the dorsolateral prefrontal cortex appear to be related to the distinctive organization of the mediodorsal thalamic nucleus and prefrontal cortex in this species.

The organization of hypothalamocerebellar cortical fibers in the squirrel monkey (Saimiri sciureus).

The organization and distribution of hypothalamocerebellar cortical fibers in squirrel monkey were investigated by using horseradish peroxidase (HRP, WGA-HRP) and 3H-leucine as anterograde tracers. Following hypothalamic injections, anterogradely labeled fibers coursed bilaterally through the periventricular gray (ipsilateral preponderance) and into the cerebellar white matter. Sparse numbers of labeled fibers appeared to descend into the reticular formation and enter the cerebellum via the brachium pontis. The pattern of cerebellar cortical labeling does not conform to that of mossy or climbing fibers. Labeled axons enter and branch within the granular layer, proceed around Purkinje cell somata, and enter the molecular layer. Within the latter some labeled fibers branch outwardly in a fanlike manner whereas others ascend before branching. Many fibers within the molecular layer ultimately assume an orientation that is similar to that of parallel fibers. The distribution patterns of hypothalamocerebellar cortical axons resemble those reported for monoaminergic fibers in the cerebellar cortex. Afferent fibers to the cerebellar cortex (including hypothalamocerebellar) that do not terminate as mossy or climbing fibers may collectively constitute a third general category of cerebellar afferent axons. On the basis of their distribution within all cortical layers these fibers are designated as multilayered fibers. The morphology of multilayered fibers stands in contrast to the presumptive mossy fiber labeling seen in lobules IX and X following large injections. Such labeling may represent a subpopulation of hypothalamocerebellar fibers or result from enzyme deposition in areas bordering the hypothalamus that project to cerebellar structures.

Somatotopic organization in cat spinal cord segments with fused dorsal horns: caudal and thoracic levels.

We have explored the somatotopic organization of the two cat spinal cord regions where the dorsal horns are fused (i.e., continuous across the midline): the caudal and thoracic segments. We have mapped the low-threshold component of dorsal horn cell receptive fields (RFs) in these segments and have charted the locations of dorsal root low-threshold mechanoreceptive dermatomes. We also have determined the projections of caudal and thoracic dorsal roots to laminae III and IV by using degeneration techniques. The dorsal skin of the tail or thorax is represented laterally, and ventral skin is represented at the midline, in the fused dorsal horns. Many caudal and thoracic dorsal horn units had RFs that crossed the dorsal or ventral midline of the skin; these units were encountered near the edges or the midline, respectively, of the fused dorsal horns. The tail is fully represented within dorsal root dermatomes S3 to Ca5. Roots more caudal than Ca5 represent progressively smaller skin areas of the distal tail. Adjacent dermatomes overlapped 15-65%. Thoracic dermatomes had a nearly vertical orientation; adjacent dermatomes overlapped by 30-75%. Dorsal roots in caudal and thoracic regions have crossed projections to the medial and lateral (but not middle) portions of the contralateral dorsal horn. These crossed projections are a possible anatomical substrate for RFs that cross the ventral or dorsal midline. The dorsal root projection patterns are consistent with those that would be predicted from the dorsal root dermatomes and dorsal horn cell somatotopy, assuming that the presynaptic terminals' somatotopy is in register with that of dorsal horn cells (the presynaptic somatotopy hypothesis; see Ref. 12).

Morphologic evidence for fiber sorting in the fasciculus cuneatus.

This report describes the medullary course and projection patterns of cervical dorsal root nerve fibers in five mammalian species (bushbaby, tree squirrel, raccoon, potoroo, and brush-tailed possum). After cutting a single cervical dorsal root and allowing appropriate postoperative survival, we studied serial, Fink-Heimer impregnated sections of the medulla from each individual to locate degenerated afferent fibers and endings. Fibers from those rostral (C2) or caudal (C8) cervical roots studied traverse the medulla as a single bundle, forming collaterals en route into the nucleus cuneatus and external cuneate nucleus at all levels. Fibers from most cervical roots (C4 through C7, sometimes C3), however, separate into two discrete bundles as they project rostrally. A more medial, superficially placed group of fibers projects predominately to the dorsal part of nucleus cuneatus, which seems to be somatotopically arranged. The second fiber group, larger, deeper, and more laterally situated, projects mainly to ventral and ventrolateral nucleus cuneatus (asomatotopically) and to the external cuneate nucleus. Based on segmental and species differences in the presence of this fiber separation, and on prior physiologic and anatomic evidence, we suggest that this form of fiber sorting in the cuneate fasciculus is modality based. It appears likely that fibers in the medial group are of primarily cutaneous origin, and that many if not most of those in the larger lateral population arise from deep receptors in muscle and joints.

Cerebellar cortical efferent fibers in the North American opossum, Didelphis virginiana. I. The anterior lobe.

The distribution of cerebellar corticonuclear and corticovestibular fibers from the anterior lobe of the North American opossum, Didelphis virginiana, was studied using the Fink and Heimer ('67) technique. Corticonuclear fibers from medial areas of anterior lobe project into the medial cerebellar nucleus (NM) in a topographically organized manner. Fibers from lobules II and III enter rostral and rostrodorsal NM, while those from lobules IV and V terminate in progressively more caudal parts of the nucleus. Collectively the terminal fields in NM for axons from lobules II-V occupy about the rostral two-thirds of the nucleus. Those areas of lobules II-V of opossum that project into NM presumably correspond to zone A of cat and primate. Cerebellar corticovestibular fibers originate from cortex located immediately lateral to areas projecting to NM. The predominance of corticovestibular projections into the lateral vestibular nucleus suggests the presence of a B zone and identifies its points of interface with the A zone. The results further suggest that zones A and B overlap at their respective margins. In contrast to other mammals studied to date, zones A and B of opossum anterior lobe are comparatively wide. Corticonuclear fibers to anterior and posterior interposed nuclei and to the lateral cerebellar nucleus (NL) originate from relatively narrow lateral portions of anterior lobe. These results also suggest that the intermediate cortex of opossum anterior lobe is not clearly divisible into individual zones C1,C2 or C3. The cortical area that innervates NL is very narrow and presumably corresponds to zone D of other forms.

Cerebellar cortical efferent fibers in the North American opossum, Didelphis virginiana. II. The posterior vermis.

The projection pattern of corticonuclear and corticovestibular fibers from vermis lobules VI-X of the North American opossum, Didelphis virginiana, was studied using a modification of the Fink and Heimer ('67) technique. The evidence suggests that corticovestibular projections in opossum are ipsilateral and exit the cerebellum via the juxtarestiform body. Lobules VI and VII contribute few, if any, corticovestibular projections. Corticovestibular fibers from lobule VIII are sparse to moderate and those from lobules IX and X are numerous. The main targets of corticovestibular fibers are the superior and spinal vestibular nuclei with some input to the medial vestibular nucleus, particularly from lobules IX and X. Although degenerated axons course through the lateral vestibular nucleus, they do not appear to terminate therein. Corticonuclear fibers from lobules VI-X are ipsilateral and terminate in the caudal and caudoventral one-third of the medial cerebellar nucleus (NM). Fibers from lateral areas of some vermal lobules appear to enter contiguous parts of the immediately adjacent posterior interposed nucleus. Although each lobule projects into a specific area of caudal and caudoventral NM when the terminal fields for lobules VI-X are superimposed, it is apparent that they are largely coextensive. This is in contrast to the pattern seen in projections from the anterior vermis. The results further indicate that zones A and B are present in lobules VI-X of opossum.

Projections of hindlimb dorsal roots to lumbosacral spinal cord of cat.

Single dorsal roots of spinal nerves that contribute to the cat lumbosacral plexus (L3-S2) were cut to evoke degeneration of centrally projecting axons. Serial sections throughout lumbosacral cord levels were impregnated by the Fink-Heimer method (20) to permit charting of the distribution patterns of segmental dorsal root afferent fibers. Afferent fibers that enter a single dorsal root have an extensive distribution to multiple cord segments; their longitudinal extent varies with entry level and with laminar targets. Afferent projections to the ventral horn reach motor nuclei only in their entry segment and the adjacent segments just above and below their entry. Those afferent fibers projecting to intermediate gray (laminae VI and VII) have the most extensive spinal distribution of any types; they may, from a single dorsal root, reach as many as 13 or 14 cord segments. Dorsal horn projections of single roots are also longitudinally expansive. Small-diameter afferent fibers course rostrally and caudally in Lissauer's tract (LT) for up to 9-10 segments. They appear to terminate in at least laminae I and II in and near their entry segment; their endings are difficult to demonstrate at greater distances where they are probably less dense. Larger caliber axons entering the dorsal horn generate a somatotopically organized projection, especially to laminae III and IV. Collaterals of these fibers appear to course longitudinally within the gray matter and they distribute to as many as six to seven segments.

Observations on the early development of ascending spinal pathways. Studies using the North American opossum.

The development of ascending spinal pathways has been studied in the North American opossum using degeneration methods and the retrograde transport of horseradish peroxidase. Axons from caudal thoracic and/or lumbosacral levels of the spinal cord reach the lateral reticular nucleus, the inferior olivary complex, the reticular formation of the medulla and pons as well as the cerebellum very early in development. Innervation of the nucleus gracilis occurs somewhat later. Spinal axons grow into most of the caudal brain stem areas they occupy in the adult animal, including the nucleus gracilis, before there is convincing evidence that they reach the thalamus. Although spinal axons enter the cerebellum early in development their adult distribution with its characteristic discontinuities appears relatively late.

Bilateral secretory responses of the adrenal medulla during stimulation of hypothalamic or mesencephalic sites.

Bilateral secretion of epinephrine and/or norepinephrine from adrenal medullae occurred in 76% of cats studied following unilateral electrical stimulation of sites in the hypothalamus or rostral mesencephalon. Responses were generally greater on the ipsilateral side. In most experiments, epinephrine alone or both epinephrine and norepinephrine outputs were increased. In only two experiments norepinephrine output was increased. Stimulation of most hypothalamic sites that evoked bilateral secretion was accompanied by an increase in blood pressure. Stimulation of effective sites in the mesencephalon was not generally accompanied by blood pressure changes.

Neurons of the basolateral amygdala: a Golgi study in the opossum (Didelphis virginiana).

The cytoarchitecture of the opossum basolateral amygdala was studied using Golgi techniques. The neuronal morphology was similar in all nuclei of the basolateral complex, an three distinct cell classes were recognized. Class I neurons, which vary in size in different nuclei, have spiny dendrites and long, projection axons. Axon hillocks and initial axonal segments often have spinous protrusions, while more distal portions of the axon give off several beaded collaterals that arborize primarily in the vicinity of the cell. Class II neurons are smaller, spine-sparse cells that are found in all nuclei of the basolateral amygdala but are greatly outnumbered by class I neurons. Axons branch and give off beaded collaterals which form a moderate to dense arborization within the dendritic field of the cell. Class II neurons exhibit considerable morphologic variability including one subtype that resembles the chandelier cell of the cerebral cortex. Varicosities (1.0 - 1.5 micrometers swellings) found along the axonal collaterals of these amygdaloid chandelier cells do not have a uniform distribution but tend to be aggregated. Segments of the collaterals displaying such clustered varicosities sometimes form nest-like entanglements. Clusters of varicosities have been observed forming multiple contacts with initial segments of class I axons. Class III neurons are neurogliaform cells which have many short, varicose dendritic branches that contact dendrites of class I neurons. Only the initial portions of their axons were impregnated. This study indicates that many of the cell types seen in the generalized, metatherian opossum are similar to those described in more specialized, placental mammals. This is the first description of amygdaloid chandelier cells and their contacts with the spiny initial segments of class I projection neurons.