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Organization of the efferent projections of the medial superior olivary nucleus in the cat as revealed by HRP and autoradiographic tracing methods.

Features of the organization of the efferent axonal projections from the medial superior olivary nucleus (MSO) in the cat were studied. In order to determine the origin and distribution of projections from MSO, the retrograde horseradish peroxidase (HRP) and autoradiographic tracing methods were used. The results showed that (1) in both HRP and autoradiographic studies the projection to the inferior colliculus was largely ipsilateral, although a contralateral component was present; (2) the projection field of MSO was confined to the ventral division of the central nucleus of the inferior colliculus, and within this field the labeling was heavier in the rostral and dorsolateral parts of the ventral division; (3) the projection to the inferior colliculus was topographic with ventral parts of MSO projecting ventrally and dorsal parts of MSO projecting dorsolaterally; (4) the projection field in the central nucleus formed successive laminae oriented from ventrolateral to dorsomedial; (5) the axonal course was via the medial or internal segment of the lateral lemniscus; and (6) some fibers in this course ended additionally within the dorsal nucleus of the lateral lemniscus. This latter projection was also topographically organized. These observations supported previously described features of lamination and tonotopic order for afferents of the inferior colliculus, as well as recent suggestions that functional segregation of afferent connections exists within the laminated portion of the central nucleus of the inferior colliculus.

Animals↗

The lateral suprasylvian corticotectal projection in cats.

The projection from the lateral suprasylvian visual areas to the superior colliculus was investigated in cats using both anterograde and retrograde tracing techniques. The retrograde transport of horseradish peroxidase (HRP) or wheat germ agglutinin-HRP (WGA-HRP) from their site of deposit in the superior colliculus indicates that all divisions of the lateral suprasylvian visual areas project to both the superficial and deep layers of the superior colliculus. However, following tracer deposits in the superior colliculus that are confined to the layers below the stratum opticum (deep layers), more neurons are labeled along the lateral bank than along the medial bank of the middle suprasylvian sulcus. Conversely, tracer deposits in the superior colliculus dorsal to and including the stratum opticum label more cells in the medial than the lateral bank. These retrograde experiments also confirm that the visual cortex along the lateral gyrus (areas 17 and 18) projects to the superficial, but apparently not to the deep layers. The visual area in the cortex surrounding the caudal two-thirds of the anterior ectosylvian sulcus projects to the deep, but not to the superficial layers. The laminar and areal patterns of anterograde axon labeling in the superior colliculus were examined after single deposits of 3H-amino acids (autoradiography), HRP, or WGA-HRP in the lateral suprasylvian cortical regions, or combined isotope and WGA-HRP deposits. Axon labeling in the superior colliculus is generally densest in the stratum opticum and extends either dorsally into the superficial layers or ventrally into the intermediate gray layer. Specifically, the anterior divisions of the lateral suprasylvian cortex project primarily to the lateral portion of the superior colliculus, with the projection from the medial bank biased toward the superficial layers and axons from the lateral bank aimed mainly at the intermediate gray layer with some axons even reaching the deepest gray layer of the superior colliculus. Both the posteromedial and posterolateral divisions of the lateral suprasylvian cortex project to more extensive portions of the mediolateral and rostrocaudal dimensions of the superior colliculus than the anterior divisions. However, the posterolateral division projects more heavily to the intermediate gray layer than the posteromedial division; from the latter, axons distribute more superficially in the superior colliculus. Finally, the cortex surrounding the posterior suprasylvian sulcus projects primarily to the medial part of the superficial layers of the superior colliculus.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Topographical organization of the inferior collicular projection and other connections of the ventral nucleus of the lateral lemniscus in the cat.

The topographic distribution of projections from the ventral nucleus of the lateral lemniscus (VNLL) in the cat was investigated with the autoradiographic tracing method. The origin of minor projections was verified by retrograde tracing methods. Small injections of tritiated leucine were placed in restricted zones of VNLL. A major afferent fiber system to the inferior colliculus was labeled in all cases. From the injection site labeled fibers coursed through and around the nuclei of the lateral lemniscus to enter the ipsilateral inferior colliculus. Regardless of the position or small size of the injection, labeled fibers distributed widely in the inferior colliculus. Fibers ended in the central nucleus and deeper layers of the dorsal cortex in most cases. There was also labeling in the ventrolateral nucleus, but very few fibers ended as lateral as the lateral nucleus. A small number of labeled fibers passed from the inferior colliculus into the nucleus of the brachium of the inferior colliculus and adjacent tegmental areas. Some labeled fibers entered the commissure of the inferior colliculus where they were traced into the dorsal cortex and rostral pole of the inferior colliculus on the side contralateral to the injection site. Though the projections labeled in individual cases were similar in their divergent pattern within the central nucleus of the inferior colliculus, specific variations in the pattern were found. The dorsal zone of VNLL projected more heavily to the deeper layers of the dorsal cortex and an adjacent field in the central nucleus than the other zones. Dorsal injections in the middle zone of VNLL, on the other hand, labeled the medial part of the central nucleus more heavily, whereas ventral injections in the middle zone resulted in heavier lateral labeling. The ventral zone of VNLL projected heavily to a central field in the central nucleus. In addition to this major afferent system of VNLL to the inferior colliculus, a smaller descending projection was found. The descending projection ended mainly in the dorsomedial periolivary region and ventral nucleus of the trapezoid body. However, in some cases a few fibers were traced to the cochlear nuclei. Finally, we observed projections to the medial geniculate body from the dorsal and ventral zones of VNLL that ended diffusely in the medial division of the medial geniculate body. Possibly some fibers from the dorsal zone contribute to a broader projection of the lateral tegmentum to the dorsal division of the medial geniculate body.

Animals↗

The neostriatal mosaic. I. Compartmental organization of projections from the striatum to the substantia nigra in the rat.

Combined neuroanatomical techniques were used to examine the organization of the striatal projection to the substantia nigra in the rat. Both double anterograde axonal tracing methods (Phaseolus vulgaris leuco-agglutinin (PHA-L) and 3H-amino acid tract tracing) and double fluorescent retrograde axonal transport tracing methods were used to examine the relationship among striatal neurons projecting to separate areas of the substantia nigra. Additionally, the distributions of retrogradely labeled striatonigral projection neurons were charted relative to the neurochemically distinct striatal "patch" compartment, identified by substance P- or leu-enkephalin-like immunoreactivity, and the complementary "matrix" compartment, identified by somatostatin-like immunoreactive fibers. These studies show two distinct types of organization in the striatonigral projections. One type is topographic in that the mediolateral relationships among these striatal efferent neurons are roughly maintained by their termination patterns in the substantia nigra, while the dorsoventral relationships are inverted. Projections from any part of the striatum, however, are distributed throughout the rostrocaudal axis of the substantia nigra. Despite their general topographic organization, the variable and dispersed nature of such projections from individual striatal loci results in partial overlap of afferent fields from separate striatal areas. The second type of organization is nontopographic and provides a different system for convergence of inputs from separated striatal areas that is superimposed on the rough topographic system. In this other projection system the mediolateral and dorsoventral relationships typical of the topographically ordered system are not maintained and are sometimes reversed. For example, PHA-L injected into the dorsal striatum labels a topographic (inverted relationship) projection to the ventral substantia nigra pars reticulata but also a smaller and separate projection to the dorsal pars reticulata and adjacent pars compacta. Retrograde tracer deposits in the pars compacta label neurons in the ventral striatum (the inverted relationship) but also clusters of neurons in the dorsal striatum. These clusters are in the neurochemically defined patch compartment whereas neurons in the matrix are labeled by injections into the pars reticulata. The dendrites of both retrogradely filled patch and matrix neurons are confined to the compartment containing their cell bodies, suggesting a restriction that would functionally segregate extrinsic striatal afferents shown in other studies to be confined to either patches or matrix.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

The organization of divergent axonal projections from the midbrain raphe nuclei in the rat.

The intranuclear organization of divergently projecting neurons of the midbrain raphe in the rat was studied by using double retrograde axonal tracing. Paired injections of the tracers N-[acetyl-3H] WGA and horseradish peroxidase were made within known projection targets of the midbrain raphe (caudate-putamen, amygdala, hippocampus, substantia nigra, and locus coeruleus). After injections of either tracer in the aforementioned targets, retrograde labeled neurons were found mainly ipsilaterally and within midline portions of the dorsal raphe nucleus, its caudal B6 portion, and within the linear and superior central nuclei of the median raphe complex. There are discrete intranuclear distributions of raphe neurons that project to these forebrain and brainstem sites, and there is an overall rostrocaudal topographic order within the raphe with neurons projecting to the neostriatum, amygdala, and substantia nigra residing most rostrally and neurons projecting to the hippocampus and/or locus coeruleus occupying caudal portions of the B6 and superior central nuclei. Such distributions of projection neurons suggest the existence of an "encephalotopic" intranuclear organization within the raphe; that is, each central nervous system structure that receives midbrain raphe projections has its own unique representation within a topographically distinct portion of one or more of the raphe subgroups. These findings suggest an overall functional organization within the midbrain raphe nuclear complex whereby rostral portions are associated with the basal ganglia and related nuclei, and caudal portions relate to the limbic system. An intermediate representation of amygdala-projecting raphe neurons functionally conjoins the two. Collateralized neurons are found within complex zones of overlap in the topographically organized distributions of raphe neurons projecting to functionally related structures.

Amygdala↗

The origin, course, and termination of the hippocampothalamic projections in the macaque.

The projections from the hippocampal formation to the thalamus were investigated with both anterograde and retrograde tracers. Horseradish peroxidase was injected into medial and midline thalamic sites in six cases, and tritiated amino acids were injected into the hippocampal formation in nine others, five of which had prior transections of the fornix. Only the subicular and entorhinal cortices were found to project to the thalamus. From the subicular cortex, dense bilateral projections were traced through the fornix to the anterior nuclei, while lighter fornical projections terminated in other rostral midline sites, including the nuclei reuniens, centralis latocellularis, and paraventricularis. These projections arose predominantly from the polymorphic cells which are located in the deepest cellular layers of the subiculum and prosubiculum. In addition, the subicular cortex was found to project to the nucleus lateralis dorsalis. The latter projection, which showed evidence of a crude topographic organization, ran either through the fornix or, unlike the other subicular efferents, through the sublenticular limb of the internal capsule to form part of the temporopulvinar bundle of Arnold. The nonfornical projection to the nucleus lateralis dorsalis passed through the medial pulvinar, where there was some additional termination. Few, if any, projections from the entorhinal cortex to the thalamus travelled in the fornix. Rather, the entorhinal efferents were carried in the inferior thalamic peduncle to the magnocellular portion of the nucleus medialis dorsalis, and in the internal capsule and bundle of Arnold to the medial pulvinar and the nucleus lateralis dorsalis.

Afferent Pathways↗

Multiple cortical targets of one thalamic nucleus: the projections of the ventral medial nucleus in the cat studied with retrograde tracers.

The organization of the cortical projections of the ventral medial thalamic nucleus (VM) was studied in the cat with retrograde tracers. The extent of the VM-cortical projections was first investigated with horseradish peroxidase injected in different cortical fields. The results obtained in the experiments indicated that the main target of VM efferents is represented by a large territory anterior to the cruciate sulcus involving area 6 and the gyrus proreus and extending into the anterior part of the medial cortical surface. The afferents to these precruciate fields arise from throughout the VM. In addition, the lateral third of VM projects upon the lateral precruciate cortex that is coextensive with the precruciate part of area 4, whereas VM efferents do not extend into the posterior sigmoid gyrus. A second major target of VM efferents is represented by the insular cortex in the anterior sylvian gyrus. VM projections also reach the prepyriform cortex and the cingulate gyrus. An anteroposterior decrease of density was found in the VM-cingulate projections. Sparse VM projections reach the temporal cortex, the adjacent posterior sylvian and ectosylvian fields, and the anterior ectosylvian gyrus. No VM projections were found either upon the visual areas 17 and 18 or upon the primary auditory cortex. The interrelations between some VM-cortical cell populations and their divergent collateralization were studied by using double retrograde labeling with fluorescent tracers. The results of these experiments demonstrated that a relatively high number (at least 20%) of VM cells projecting to the insula are also connected to the precruciate fields by means of axon collaterals. This finding indicates that VM is a highly collateralized structure of the cat's thalamus. Very few branched cells were found in the other combinations of cortical fields here examined (precruciate vs. posterior sylvian fields, lateral precruciate vs. proreal cortex, anterior vs. posterior cingulate fields). Altogether these data indicate that VM branched cells preferentially interconnect the two main cortical targets of the nucleus, i.e., precruciate and insular fields. The results of the present study are discussed in regard to the literature on the VM projections in the rat and the previously available data in the cat, to the afferent VM organization in the cat, to the relationships between VM and the nucleus submedius, and to the anatomical and functional role of VM in relation to the so-called "nonspecific" thalamocortical system.

Animals↗

The ipsilateral retinocollicular projection in the rabbit: an autoradiographic study of postnatal development and effects of unilateral enucleation.

The postnatal development of the ipsilateral retinocollicular projection in the rabbit and the effects of unilateral enucleation (performed on the day of birth, day 0) on that development were studied by using the anterograde axonal transport of tritiated proline injected intraocularly. Material from 1-, 6-, and 10-day-old (i.e., at days 1, 6, and 10) and adult animals was examined. On day 1, autoradiographically labelled optic fibers from the ipsilateral eye formed distinct patches and bands within the superior colliculus (SC), which were restricted primarily to the lateral one-half and anterior one-third to one-half of the nucleus. At subsequent ages no major changes in the location of this projection were found for normal animals or animals enucleated on day 0 (0-DE animals). From dorsal-view reconstructions, the pattern of the ipsilateral projection appeared wedge-shaped with a broad base aligned with the lateral SC border for all normal and 0-DE animals at the various postnatal ages examined. In normal animals the surface area of this projection increased with age and maintained a constant proportion of the increasing surface area of the total SC. In 0-DE animals the surface area of the projection initially increased more rapidly than in normal animals. Thus, by day 6 the area was already within the normal adult range but did not exceed this range at later postnatal ages. The only obvious difference in the appearance of the ipsilateral retinocollicular projection between normal and 0-DE animals at corresponding ages was an enhanced radial distribution of the projection across laminae in the 0-DE animals. Taken together these findings suggest that, in the rabbit, once topographically appropriate connections are established between the SC and the ipsilateral retinal projection, they are maintained regardless of substantial postnatal growth of the SC and removal of the contralateral retinal projection to the SC.

Age Factors↗

Organization of amygdaloid projections to the mediodorsal thalamus and prefrontal cortex: a fluorescence retrograde transport study in the rat.

Previous studies have shown that the amygdala projects to both the mediodorsal thalamic nucleus (MD) and its cortical projection area, the prefrontal cortex (PFC). In this investigation rats received injections of different fluorescent retrograde tracers (true blue and diamidino yellow) into MD and either the lateral, polar, or medial PFC in order to examine the relationship of amygdaloid neurons with cortical and/or thalamic projections. PFC injections labeled neurons in the basolateral (BL), basomedial (BM), ventral endopiriform (EnV), and rostral lateral nuclei as well as the periamygdaloid cortex (PAC) and the medial part of the amygdalohippocampal area (AHA). In BL, which contained the great majority of neurons projecting to PFC, most labeled cells were concentrated in particular parts of the nucleus and were topographically organized. The overwhelming majority of labeled neurons in BL were large pyramidal or piriform cells that correspond to class I neurons described in Golgi studies. Occasional small neurons with thin dendrites were also observed; these cells may be class II neurons. MD injections labeled numerous cells in the anterior division of the cortical nucleus, medial nucleus, and caudomedial part of the central nucleus. Moderate numbers of labeled cells were found in caudal portions of BM and PAC, whereas scattered cells were observed throughout the rest of the amygdala with the exception of the lateral nucleus. In BL and AHA many MD-projecting neurons were observed along nuclear boundaries and in the adjacent white matter. Neurons in BL, BM, and AHA usually had large elongated or irregular somata and two to four primary dendrites that branched sparingly. Other cells had smaller ovoid somata. The morphology and distribution of MD-projection cells in the basolateral amygdala indicate that they are primarily large class II neurons. Double-labeled amygdaloid neurons, labeled by both cortical and thalamic injections, were observed only in a small number of animals. Control experiments suggest that most of the double-labeled cells in these cases were artifacts caused by spread of the thalamic injectate into the third ventricle with subsequent uptake by fibers in the anterior commissure. Thus the findings of this study suggest that different neuronal populations in the amygdala project to the two poles of the MD-PFC system. In the basolateral amygdala class I neurons are the predominant cell type involved in PFC projections, whereas a subpopulation of class II neurons, hitherto thought to be primarily local-circuit neurons, project to MD.

Amidines↗

Origin of ascending projections to an isofrequency region of the mustache bat's inferior colliculus.

The inferior colliculus of the mustache bat is similar in most respects to the inferior colliculus of more commonly studied mammals, but one isofrequency contour, the dorsoposterior division, is greatly overrepresented. The present study utilizes this specialization of the auditory system in the mustache bat to determine the total set of ascending projections to a single isofrequency contour of the inferior colliculus. Within the dorsoposterior division, neurons are all very narrowly tuned to 60 kHz, the major component of this bat's echolocation call. The afferent projections to this isofrequency contour were identified by making deposits of horseradish peroxidase (HRP) within the dorsoposterior division after physiologically defining its borders. Two other frequency representations are present in the central nucleus of the inferior colliculus of the mustache bat, the anterolateral division in which there is an orderly progression of frequencies from 59 down to 20 kHz, and the medial division in which frequencies from 63-120 kHz are represented. In additional experiments, the afferent projections to the medial and anterolateral divisions were examined, providing an anatomical description of the tonotopicity of the lower auditory nuclei. Deposits of HRP in the DPD labeled cells in each of the lower brainstem auditory nuclei that have previously been shown to project to the entire central nucleus of the inferior colliculus. The ascending projections to the dorsoposterior division include contralateral projections from the cochlear nucleus and inferior colliculus, ipsilateral projections from the medial superior olive, ventral and intermediate nuclei of the lateral lemniscus, and bilateral projections from the lateral superior olive and dorsal nucleus of the lateral lemniscus. In most of the nuclei, labeled cells were confined to specific portions of the nuclei, often forming "slabs" of labeled cells across the rostrocaudal extent of most nuclei. These slabs presumably represent the 60 kHz representation in each of the lower nuclei. When deposits of HRP were made into other frequency band representations of the inferior colliculus, in either the medial or anterolateral division, labeled cells again formed slabs in each lower nucleus. However, the location of the slab varied as a function of the best frequency of neurons at the deposit site, and labeled cells were not present within the 60 kHz representation. These results show the general tonotopy of the mustache bat's brainstem auditory nuclei, and with respect to the dorsoposterior division, clearly reveal the total set of projections to a single isofrequency region.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Central projections of the rat radial nerve investigated with transganglionic degeneration and transganglionic transport of horseradish peroxidase.

Transganglionic degeneration and transganglionic transport of HRP were used for investigation of the spinal cord and brainstem projections from the superficial, cutaneous (SR) and deep, muscular (DR) branches of the radial nerve. The HRP study included a numerical and size analysis of labelled dorsal root ganglion (DRG) cells. In degeneration experiments the SR nerve was found to project somatotopically to laminae III-IV, but degeneration was also found in lamina I and inconsistently in lamina II. Transection of the DR nerve was found to give rise to a small amount of degeneration, which in "sham" operations was established to result from the skin injury during dissection of the DR nerve. With the HRP method, the SR nerve was found to project somatotopically to laminae I-IV, whereas the DR nerve projected more diffusely to the medial part of laminae V-VII. HRP application to the SR and DR nerves resulted in labelling of a mean of 1,024 and 310 DRG cells, respectively. These labelled neurons had a median cell area of 381 and 562 micron 2 for the SR and DR nerves, respectively, and both small and large cells were labelled in both types of experiments. In the lower brainstem, projections from the SR nerve were found only in the ipsilateral dorsal part of the main cuneate nucleus (MCN) with both methods. Brainstem projections from the DR nerve that were found only with the HRP method were found in the ipsilateral ventral part of the MCN together with a projection to the ipsilateral external cuneate nucleus. No projections were found to the central cervical nucleus. The present results indicate that cutaneous compared to muscular primary sensory neurons are much more prone to react with transganglionic degeneration after peripheral nerve transection. Furthermore, in the rat the SR nerve projects somatotopically, whereas the DR nerve does not. Both nerve branches are connected to small and large spinal ganglion cells, although the median cell area is larger in muscular neurons.

Afferent Pathways↗

Collateralization of the amygdaloid projections of the rat prelimbic and infralimbic cortices.

Previous studies indicate that the distribution of corticoamygdaloid neurons in the rat prelimbic (PL) and infralimbic (IL) cortices overlaps with the distribution of neurons projecting to the contralateral medial prefrontal cortex (MPC), insular cortex, mediodorsal thalamus, and dorsal medulla. In view of the poorly differentiated cytoarchitecture of PL and IL, and their designation as cortical regions transitional between the allocortex and isocortex, the present study sought to determine whether several cortical and subcortical projections from these areas arise as collaterals of corticoamygdaloid neurons. Injections of the fluorescent dyes Fast Blue (FB) or bisbenzimide (BB) were made into the amygdaloid complex and the following areas: agranular and granular insular cortices; mediodorsal thalamic nucleus (MD); nucleus tractus solitarii/dorsal medulla (NTS); contralateral amygdaloid complex; and ipsilateral and contralateral MPC. Neurons projecting to the ipsilateral amygdaloid complex were located mainly in layers II and V with fewer cells in layer III. Concomitant injections into the insular cortex, MD, and NTS labeled populations of neurons arranged in laminae that partially overlapped with, but were essentially separate from, corticoamygdaloid neurons. Projections to the insular cortex arose from layers II and V; those to MD arose from layers V and VI. Corticobulbar projections from IL originated from neurons arranged in a thin lamina in the deep part of layer V. Very few neurons projecting to both the amygdaloid complex and any of these areas were observed. Bilateral injections of FB and BB into the amygdaloid complex producted very few double-labeled cells in PL and IL. Further, in layer V, ipsilaterally projecting corticoamygdaloid neurons tended to be located more deeply than contralaterally projecting neurons. Combined injections of BB and FB into the amygdaloid complex and the contralateral (but not ipsilateral) MPC resulted in significant numbers of double-labeled neurons in layers II, III, and V of PL and IL. Control injections of fluorescent dyes into the cerebrospinal fluid labeled few neurons in the superficial layers of PL and IL and a combined injection into the amygdaloid complex (FB) and subarachnoid space (BB) resulted in a very small number of double-labeled cells in layer II only. The results suggest that a significant proportion of neurons in PL and IL projecting to the amygdaloid complex issue collaterals innervating the contralateral MPC. Evidence is discussed that suggests that the interhemispheric collaterals of MPC corticoamygdaloid neurons may serve to correlate the amygdaloid outputs of the MPC bilaterally.

Amygdala↗

Convergence and divergence in the afferent projections to cat area 17.

We have examined the topography of the afferent connections to area 17 in the cat by means of double retrograde label tracing techniques. Injections of two fluorescent retrograde tracers, diamidino yellow and fast blue, were made with variable separations in area 17 and the spatial distributions of the resulting populations of labeled cells examined in afferent cortical areas and subcortical structures. When injections were separated rostrocaudally, the topographic organizations of the projections were characterized quantitatively with two graphic methods: the labeling density curve and the connectivity graph. The labeling density curve measures labeled neuron density in successive rostrocaudal sections, whereas the connectivity graph provides a two-dimensional model of the topography of a given connectivity. The connectivity graph makes it possible to define two parameters that characterize the topography of the connection: the convergence and the divergence. The convergence is defined as the extent of an afferent structure that contains neurons converging on a line normal to the cortical surface in area 17. The divergence is the extent of area 17 that is innervated by neurons contained in an infinitely small region of the afferent structure. The results show that a number of subcortical structures project to area 17 in a nontopographic manner, i.e., that in each of these structures neurons contained in an infinitely small region send projections to the whole of area 17 and that a line normal to the surface of area 17 is innervated by neurons distributed throughout the afferent structure in question. Nontopographic projections are found from the intralaminar nuclei, the ventral mesencephalic tegmental region, the diagonal band of Broca, and the locus coeruleus. All remaining subcortical structures and cortical areas send topographically organized projections to area 17. The extent of the convergence and divergence, however, varies between structures. Only the projection from the A laminae of the LGN was found to approximate a point-to-point projection with a convergence of 0.4 mm and 2 mm in divergence. Much larger convergence and divergence values are found in the projections from the claustrum and the cortical areas. For example, the divergence reaches 20 mm for the projections from area 20 or from the anterior part of the lateral suprasylvian sulcus. Knowing the convergence and divergence values and the retinotopic organizations of area 17 and a number of its afferents, it becomes possible to test whether connections in the visual system link regions representing the same zone of the visual field.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Anatomical mapping of the cerebellar nucleocortical projections in the rat: a retrograde labeling study.

An analysis of the cerebellar nucleocortical projections was made by means of retrograde cellular labeling with wheat germ agglutinin-horseradish peroxidase conjugate. Each of the main nuclear subregions appears to give rise to nucleocortical projections. The cortical distribution of the projections is referred to here in term of sagittal zones. Zones A, B, and C conform to the recent description in the rat (Buisseret-Delmas, '88a,b) on the basis of their olivocortical and corticonuclear projections. A corresponding description of zone D is given here. According to their distribution, three types of nucleocortical projections have been distinguished: 1) ipsilateral, reciprocal; 2) nonreciprocal; and 3) contralateral, symmetrical to the corticonuclear afferent. Reciprocal projections are strictly arranged in the sagittal direction, with the following zonal distribution. Zone A is subdivided into two subzones. Medial A zone receives its nuclear afferents from the medial aspect of the nucleus medialis (NM). The lateral A zone of the anterior lobe and lobule VI and that of the posterior lobe receive their reciprocal nuclear afferents from the ventrolateral NM and the dorsolateral protuberance, respectively. Zone B does not seem to receive nucleocortical projections. Zone C has three subzones in the rat. C1 is supplied from the medial third of the anterior and posterior subdivisions of the nucleus interpositus (NIA and NIP, respectively). C2 is supplied from the central third of the NIA and NIP. Rostrocaudally, the anterior lobe and lobule VIII are connected to the NIA, and lobules VI and VII to the NIP. C3 appears to be connected to the lateral third of NIA. Zone D contains three subzones mediolaterally in the rat. D0, not previously described, is defined on the basis of both its olivary afferent from the medial half of the ventral lamella of the principal olive and its corticonuclear projections onto the dorsolateral hump of Goodman et al. ('63). It receives a reciprocal nucleocortical afferent from the dorsolateral hump. D1 receives its olivary afferent from the dorsal lamella of the principal olive. It is reciprocally connected with the lateral, magnocellular part of the nucleus lateralis (NL). D2 is the most lateral subzone of the hemisphere. Its olivary afferent comes from the lateral half of the ventral lamella of the principal olive. D2 is reciprocally connected with the ventral, parvicellular subdivision of NL. The main cortical recipients for the nonreciprocal projections are the lateral A zone, the C3, and the D1 subzones.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Cells of origin of spinothalamic tract projections to the medial and lateral thalamus in the cat.

The double fluorescent retrograde labeling method was used to examine the distribution of spinothalamic tract (STT) cells that project to the medial and lateral thalamus in the cat. Injections of one fluorescent tracer (Fast Blue or Diamidino Yellow) were made throughout the lateral thalamus and injections of the other tracer were made in the medial thalamus at sites extrapolated from recording track coordinates. Survival times were successively extended (up to 5 weeks) in order to maximize labeling in both the cervical and lumbosacral spinal cord. On average, over 2,000 labeled contralateral STT cells were counted in serial sections from segments C5-7 and L5-S2. Numerical variability of the order of a factor of two was attributable to inherent differences between individual animals. The total number of cells labeled with fluorescent tracers was comparable to the number labeled with horseradish peroxidase in control cases, although there were significant differences between the laminar distributions of labeling produced by the two methods. Injections made anterior to the thalamus to control for labeling due to leakage or passing fibers did not produce substantial spinal labeling. The laminar distribution of fluorescent dye-labeled STT cells was consistent; about half (47%) were located in lamina I, 8% were in lamina V, 5% in lamina VI, 20% in lamina VII, and 20% in lamina VIII. The proportions of STT cells in laminae I and V were higher in cervical segments (57% and 12%, respectively) than in lumbosacral segments (38% and 6%). The dominant contribution of lamina I cells to the STT thus revealed by the fluorescent tracers is striking. The proportions of STT cells labeled from the medial and the lateral thalamus varied with segmental and laminar location and with injection placement. The majority (62%) of STT cells in most cases projected only to the medial thalamus, 25% projected only to the lateral thalamus, and 13% projected to both. The STT cell populations in laminae I, VII, and VIII each displayed this common projection pattern. In contrast, cells in laminae V and VI projected predominantly to the lateral thalamus. Twice as many STT cells in lamina I (19%) projected to both the medial and the lateral thalamus as from other laminae. A greater proportion of laminae V-VIII STT cells in segments L5-6 projected to the lateral thalamus, and in S1-2, more projected to the medial thalamus.(ABSTRACT TRUNCATED AT 400 WORDS)

Amidines↗

Central projections of C4-C8 dorsal root ganglia in the rat studied by anterograde transport of WGA-HRP.

Injections of WGA-HRP were made in the rat C4-C8 dorsal root ganglia (DRGs) individually to study the central projections and their relations to each other. The main dorsal horn projections from these DRGs to the dorsal horn lamina II extended for about two segments rostrally and caudally to the injected DRG, whereas the projections to laminae I, III, and IV were less restricted rostrocaudally. Comparisons of the dorsal horn projections from the DRGs investigated indicated a tendency for a somatotopic organization, which was most prominent in lamina II. Labeled central branches from the C4-8 DRGs could be traced in the dorsal column as far caudally as 12-17 segments caudal to the level of entrance. Most of these fibers appeared to end in the medial dorsal horn base, including the column of Clarke. Labeling of primary afferents in the ventral horn generally extended for at least 3-4 segments rostral and caudal to the level of the injected DRG. Projections to the central cervical nucleus were most prominent from the C4 DRG and gradually became less prominent from the more caudal DRGs. Heavy projections to the cuneate nucleus (Cun) originated from the C7 and C8 DRG, whereas those from the C4-C6 DRGs were less extensive. The Cun projections from the different DRGs appeared to overlap, and the same was true for the projections to the external cuneate nucleus. Projections to the gracile nucleus, the vestibular nuclear complex, including nucleus X, and to trigeminal sensory nuclei were seen from all DRGs investigated.

Afferent Pathways↗

Projections from the amygdala to basoventral and mediodorsal prefrontal regions in the rhesus monkey.

The sources of ipsilateral projections from the amygdala to basoventral and mediodorsal prefrontal cortices were studied with retrograde tracers (horseradish peroxidase or fluorescent dyes) in 13 rhesus monkeys. The basoventral regions injected with tracers included the orbital periallocortex and proisocortex, orbital areas 13, 11, and 12, lateral area 12, and ventral area 46. The mediodorsal regions included portions of medial areas 25, 32, 14, and dorsal area 8. The above sites represent areas within two architectonic series of cortices referred to as basoventral or mediodorsal on the basis of their anatomic location. Each series consists of areas that show a gradual increase in the number of layers and their delineation in a direction from the caudal orbital and medial limbic cortices, which have an incipient laminar organization, towards the eulaminated periarcuate cortices (Barbas and Pandya, J. Comp. Neurol. 286: 353-375, '89). Labeled neurons projecting to the prefrontal cortex were found in the basolateral, basomedial (also known as accessory basal), lateral, and ventral cortical nuclei, and in the anterior amygdaloid and amygdalopiriform areas. The distribution of labeled neurons differed both quantitatively and qualitatively depending on whether the injection sites were in basoventral or mediodorsal prefrontal cortices. Cases with caudal orbital injections had the most labeled neurons in the amygdala, followed by cases with injections in cortices situated medioventrally. The latter received a high proportion of their amygdaloid projections from the basomedial nucleus. The lateral amygdaloid nucleus sent a robust projection to the least architectonically differentiated orbital periallocortex, and a weaker projection to the adjoining orbital proisocortical regions, but did not appear to project to either medial proisocortical sites or to the more differentiated ventrolateral or dorsolateral prefrontal cortices. In addition, there were topographical differences in the origin of projections from one amygdaloid nucleus directed to various prefrontal cortices. These differences were correlated either with the destination of the axons of afferent amygdaloid neurons to basoventral or to mediodorsal prefrontal cortices and/or with their projection to areas with varying degrees of laminar organization within the basoventral or mediodorsal sector. The clearest topography was observed for projections originating in the basolateral nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

Amygdala↗

Morphological types of projection neurons in layer 5 of cat visual cortex.

Pyramidal cells in layer 5 of the visual cortex have multiple cortical and subcortical projection sites. Previous studies found that many cells possess bifurcating axons and innervate more than one cortical or subcortical target, but cells projecting to both cortical and subcortical targets were not observed. The present study examines the morphology of cells in cat visual cortex projecting to the superior colliculus, the main subcortical target of layer 5, and cells in layer 5 projecting to cortical areas 18 and 19. The neurons that give rise to these different projections were retrogradely labelled and intracellularly stained in living brain slices. Our results show that cells within each projection group have several morphological features in common. All corticotectal cells have a long apical dendrite forming a large terminal tuft in layer 1. Their cell bodies are medium sized to large, and their basal dendrites form a dense and symmetrical dendritic field. Corticocortical cells in layer 5 have a very different morphology: their apical dendrites are short and they never reach higher than layers 2/3. Their cells bodies are small to medium sized and they have fewer basal dendrites than corticotectal cells. Thus there are two morphologically distinct projection systems in layer 5, one projecting to cortical and the other one to subcortical targets, suggesting that these two systems transmit different information from the visual cortex. Among the corticotectal cells with the largest cell bodies we found some cells whose basal and apical dendrites were almost devoid of spines. Spiny and spinefree corticotectal cells also have different intrinsic axon collaterals and therefore play different roles in the cortical circuitry. While many spiny corticotectal cells have axon collaterals that project to layer 6, spinefree corticotectal cells have fewer axon collaterals and these do not arborize in layer 6. We suggest that the two morphological types of corticotectal cells might be related to functional differences known to exist among these cells. We discuss how the presence or absence of spines affects the integration of the synaptic input and how this might be related to the cells' functional properties.

Animals↗