Showing papers in "The Journal of Comparative Neurology in 1978"

An autoradiographic analysis of the differential ascending projections of the dorsal and median raphe nuclei in the rat

Abstract: The differential projections from the dorsal raphe and median raphe nuclei of the midbrain were autoradiographically traced in the rat brain after 3H-proline micro-injections. Six ascending fiber tracts were identified, the dorsal raphe nucleus being the sole source of four tracts and sharing one with the median raphe nucleus. The tracts can be classified as those lying within the medial forebrain bundle (dorsal raphe forebrain tract and the median raphe forebrain tract) and those lying entirely outside (dorsal raphe arcuate tract, dorsal raphe periventricular tract, dorsal raphe cortical tract, and raphe medial tract). The dorsal raphe forebrain tract lies in the ventrolateral aspect of the medial forebrain bundle (MFB) and projects mainly to lateral forebrain areas (e.g., basal ganglion, amygdala, and the pyriform cortex). The median raphe forebrain tract lies in the ventromedial aspect of the MFB and projects to medial forebrain areas (e.g., cingulate cortex, medial septum, and hippocampus). The dorsal raphe cortical tract lies ventrolaterally to the medial longitudinal fasciculus and projects to the caudate-putamen and the parieto-temporal cortex. The dorsal raphe periventricular tract lies immediately below the midbrain aqueduct and projects rostrally to the periventricular region of the thalamus and hypothalamus. The dorsal raphe arcuate tract curves laterally from the dorsal raphe nucleus to reach the ventrolateral edge of the midbrain and projects to ventrolateral geniculate body nuclei and the hypothalamic suprachiasmatic nuclei. Finally, the raphe medial tract receives fibers from both the median and dorsal raphe nuclei and runs ventrally between the fasciculus retroflexus and projects to the interpeduncular nucleus and the midline mammillary body. Further studies were done to test whether the fiber tracts travelling in the MFB contained 5-HT. Unilateral (left) injections of 5,7-dihydroxytryptamine (5 μgm/400 nl) 18 days before midbrain raphe microinjections of 3H-proline produced a reduction in the grain concentrations in all the ascending fibers within the MFB. Furthermore, pharmacological and behavioural evidence was obtained to show that the 5-HT system had been unilaterally damaged; these animals displayed preferential ipsilateral turning in a rotameter which was strongly reversed to contralateral turning after 5-hydroxytryptophan administration. The results show that DR and MR nuclei have numerous ascending projections whose axons contain the transmitter 5-HT. The results agree with the neuroanatomical distribution of the 5-HT system previously determined biochemically, histochemically, and neurophysiologically. The midbrain serotonin system seems to be organized by a series of fiber pathways. The fast transport rate in these fibers was found to be about 108 mm/day.

Amygdaloid projections to subcortical structures within the basal forebrain and brainstem in the rat and cat

Abstract: The efferent fiber connections of the nuclei of the amygdaloid complex with subcortical structures in the basal telencephalon, hypothalamus, midbrain, and pons have been studied in the rat and cat, using the autoradiographic method for tracing axonal connections. The cortical and thalamic projections of these nuclei have been described in previous papers (Krettek and Price, ′77b,c). Although the subcortical connections of the amygdaloid nuclei are widespread within the basal forebrain and brain stem, the projections of each nucleus have been found to be well defined, and distinct from those of the other amygdaloid nuclei. The basolateral amygdaloid nucleus projects heavily to the lateral division of the bed nucleus of the stria terminalis (BNST), to the caudal part of the substantia innominata, and to the ventral part of the corpus striatum (nucleus accumbens and ventral putamen) and the olfactory tubercle; it projects more lightly to the lateral hypothalamus. The central nucleus also projects to the lateral division of the BNST and the lateral hypothalamus, but in addition it sends fibers to the lateral part of the substantia nigra and the marginal nucleus of the brachium conjunctivum. The basomedial nucleus has projections to the ventral striatum and olfactory tubercle which are similar to those of the basolateral nucleus, but it also projects to the core of the ventromedial hypothalamic nucleus and the premammillary nucleus, and to a central zone of the BNST which overlaps the medial and lateral divisions. The medial nucleus also projects to the core of the ventromedial nucleus and the premammillary nucleus, but sends fibers to the medial division of the BNST and does not project to the ventral striatum. The posterior cortical nucleus projects to the premammillary nucleus and to the medial division of the BNST, but a projection from this nucleus to the ventromedial nucleus has not been demonstrated. Projections to the “shell” of the ventromedial nucleus have been found only from the ventral part of the subiculum and from a structure at the junction of the amygdala and the hippocampal formation, which has been termed the amygdalo-hippocampal area (AHA). The AHA also sends fibers to the medial part of the BNST and the premammillary nucleus. Virtually no subcortical projections outside the amygdala itself have been demonstrated from the lateral nucleus, or from the olfactory cortical areas around the amygdala (the anterior cortical nucleus, the periamygdaloid cortex, and the posterior prepiriform cortex). However, portions of the endopiriform nucleus deep to the prepiriform cortex project to the ventral putamen, and to the lateral hypothalamus.

Catecholamine innervation of the basal forebrain. IV. Topography of the dopamine projection to the basal forebrain and neostriatum.

Abstract: In this study the location of dopamine (DA) neuron perikarya in the rostral mesencephalon of the rat was determined using the glyoxylic acid fluorescence histochemical technique. Subsequently the topography of the projection of these mesencephalic neurons on the basal forebrain and striatum was analyzed using the anterograde transport-autoradiographic tracing method and the retrograde transport-horseradish peroxidase (HRP) technique. The results of these anatomical studies were correlated with the biochemical and histochemical studies presented in previous reports (Moore, 1978; Fallon and Moore, 1978; Fallon et al., 1978) to provide the following conclusions. The topography of the DA neuron projection on the basal forebrain and neostriatum is organized in three planes, dorsal-ventral, medial-lateral and anterior-posterior. DA cells are found almost exclusively in the substantia nigra (SN) and ventral tegmental area (VTA). Ventral cells of the SN and VTA project to the dorsal structures of the basal forebrain such as the septum, nucleus accumbens and neostriatum. The latter includes some DA cells located ventrally in the pars reticulata of the SN. Dorsal cells project to ventral structures. The medial-lateral topography is organized such that the medial sectors of the SN-VTA area project to the medial parts of nuclei in the basal forebrain and neostriatum whereas lateral sectors of the SN-VTA area project to the lateral parts of nuclei in the basal forebrain and neostriatum. An anterior-posterior topography also is evident such that anterior parts of the SN-VTA project anteriorly whereas the posterior SN-VTA projects more posteriorly in these areas. These observations are consistent with the view that the DA neurons of the SN-VTA complex form a single nuclear group with a highly topographically organized projection innervating not only deep nuclei of the telencephalon but allocortical structures as well.

Intracortical connectivity of architectonic fields in the somatic sensory, motor and parietal cortex of monkeys

Abstract: Anterograde and retrograde transport methods were used to study the corticocortical connectivity of areas 3a, 3b, 1, 2, 5, 4 and 6 of the monkey cerebral cortex. Fields were identified by cytoarchitectonic features and by thalamic connectivity in the same brains. Area 3a was identified by first recording a short latency group I afferent evoked potential. Attempts were made to analyze the data in terms of: (1) routes whereby somatic sensory input might influence the performance of motor cortex neurons; (2) possible multiple representations of the body surface in the component fields of the first somatic sensory area (SI). Apart from vertical interlaminar connections, two types of intracortical connectivity are recognized. The first, regarded as "non-specific," consists of axons spreading out in layers I, III and V-VI from all sides of an injection of isotope; these cross architectonic borders indiscrimininately. They are not unique to the regions studied. The second is formed by axons entering the white matter and re-entering other fields. In these, they terminate in layers I-IV in one or more mediolaterally oriented strips of fairly constant width (0.5--1 mm) and separated by gaps of comparable size. Though there is a broadly systematic topography in these projections, the strips are probably best regarded as representing some feature other than receptive field position. Separate representations are nevertheless implied in area 3b, in areas 1 and 2 (together), in areas 3a and 4 (together) and in area 5; with, in each case, the representations of the digits pointed at the central sulcus. Area 3b is not connected with areas 3a or 4, but projects to a combined areas 1 and 2. Area 1 is reciprocally connected with area 3a and area 2 reciprocally with area 4. The connectivity of area 3a, as conventionally identified, is such that it is probably best regarded not as an entity, but as a part of area 4. Areas identified by others as area 3a should probably be regraded as parts of area 3b. Parts of area 5 that should be more properly considered as area 2, and other parts that receive thalamic input not from the ventrobasal complex but from the lateral nuclear complex and anterior pulvinar, are also interconnected with area 4. More posterior parts of area 5 are connected with laterally placed parts of area 6. A more medial part of area 6, the supplementary motor area, occupies a pivotal position in the sensory-motor cortex, for it receives fibers from areas 3a, 4, 1, 2 and 5 (all parts), and projects back to areas 3a, 4 and 5.

A Golgi study of cell types in the hilar region of the hippocampus in the rat.

Abstract: The morphology of neurons in the "hilar region" of the hippocampus (fields CA3c and CA4 of Lorente de No, '34) was analyzed with several variants of the Golgi technique. Hippocampi were dissected from the brains of 28-day-old rats, fixed and impregnated by immersion, and sectioned perpendicular to the long axis. Based on the resident cell types, aspects of the neuropil, and published data related to afferent termination, the area under study was divided into four zones. At least 21 cell types were observed throughout these zones, several of which had not previously been described. Many cells in this area exhibited an impressive number and variety of dendritic and axonal appendages, including spines on the proximal portion of some axons. The close apposition of fibers to these axonal spines suggested the possibility of axo-axonal interactions. The influence of dentate granule cells, through their mossy fibers, on the synaptic economy of the "hilar region" was found to be more extensive than previously reported. Mossy fibers appeared to terminate on the dendrites of several types of non-pyramidal cells, which bear no thorny excrescences, by means of thin filiform extensions which emanate from the mossy fiber expansions and by means of thin mossy fiber collaterals which are devoid of typical expansions. Consideration is given to a long-standing debate as to whether the deep "hilar region" (CA4 of Lorente de No, '34, hilus of the fascia dentata of Blackstad, '56) is related more to the hippocampus or to the fascia dentata and it is concluded that the deep hilar region is an area of mergence of the polymorphic zones of these two cortical structures. The results of the present study do not support the proposition that the deep hilar region is an extension of the pyramidal layer of the hippocampus as suggested by Lorente de No ('34), and thus CA4 is a misnomer. Rather, the cells in this area are most closely related to the fascia dentata and should thus be considered to lie in the polymorphic zone of "area dentata" as proposed initially by Blackstad ('56).

Serotonin neurons of the midbrain raphe: ascending projections.

Abstract: The ascending projections of serotonin neurons of the midbrain raphe were analyzed in the rat using the autoradiographic tracing method. Axons of raphe serotonin neurons ascend in the ventral tegmental area and enter the medial forebrain bundle. A number of fibers leave the major group to ascend along the fasciculus retroflexus. Some fibers enter the habenula but the majority turn rostrally in the internal medullary lamina of the thalamus to innervate dorsal thalamus. Two additional large projections leave the medial forebrain bundle in the hypothalamus; the ansa peduncularis-ventral amygdaloid bundle system turns laterally through the internal capsule into the striatal complex, amygdala and the external capsule to reach lateral and posterior cortex, and another system of fibers turns medially to innervate medial hypothalamus and median eminence and form a contrelateral projection via the supraoptic commissures. Rostrally the major group in the medial forebrain bundle divides into several components: fibers entering the stria medullaris to terminate in thalamus; fibers entering the stria terminalis to terminate in the amygdala; fibers traversing the fornix to the hippocampus; fibers running through septum to enter the cingulum and terminate in dorsal and medial cortex and in hippocampus; fibers entering the external capsule to innervate rostral and lateral cortex; and fibers continuing forward in the medial olfactory stria to terminate in the anterior olfactory nucleus and olfactory bulb.

Immunohistochemical localization of enkephalin in rat brain and spinal cord.

Abstract: The distribution of immunoreactive enkephalin in rat brain and spinal cord was studied by immunoperoxidase staining using antiserum to leucine-enkephalin ([Leu5]-enkephalin) or methionine-enkephalin ([Met5]-enkephalin). Immunoreactive staining for both enkephalins was similarly observed in nerve fibers, terminals and cell bodies in many regions of the central nervous system. Staining of perikarya was detected in hypophysectomized rats or colchicine pretretated rats. The regions of localization for enkephalin fibers and terminals include in the forebrain: lateral septum, central nucleus of the amygdala, area CA2 of the hippocampus, certain regions of the cortex, corpus striatum, bed nucleus of the stria terminalis, hypothalamus including median eminence, thalamus and subthalamus; in the midbrain: nucleus interpeduncularis, periaqueductal gray and reticular formation; in the hind brain: nucleus parabrachialis, locus ceruleus, nuclei raphes, nucleus cochlearis, nuclear tractus solitarii, nucleus spinalis nervi trigemini, motor nuclei of certain cranial nerves, nucleus commissuralis and formatio reticularis; and in the spinal cord the substantia gelatinosa. In contrast enkephalin cell bodies appear sparsely distributed in the telencephalon, diencephelon, mensencephalon and rhombencephalon. The results of the histochemical staining show that certain structures which positively stain for enkephalin closely correspond to the distribution of opiate receptors in the brain and thus support the concept that the endogenous opiate peptides are involved in the perception of pain and analgesia. The localization of enkephalin in the preoptic-hypothalamic region together with the presence of enkephalin perikarya in the paraventricular and supraoptic nuclei suggest a role of enkephalin in the regulation of neuroendocrine functions.

The retinotopic organization of area 17 (striate cortex) in the cat.

Abstract: The location and retinotopic organization of visual areas in the cat cortex were determined by systematically mapping visual cortex in over 100 cats. The positions of the receptive fields of single neurons or small clusters of neurons were related to the locations of the corresponding recording sites in the cortex to determine the representations of the visual field in these cortical areas. In this report, the first of a series, we describe the organization of area 17. A single representation of the cat's entire visual field corresponds closely to the cytoarchitectonically defined area 17. This area has the largest cortical surface area (380 mm2) and the highest cortical magnification factor (3.6 mm2/degree2 at area centralis) of all the cortical areas we have studied. There was perfect agreement between the borders of area 17 determined electrophysiologically and cytoarchitecturally. This area contains a first order transformation of the visual hemifield in which every adjacent point in the visual field is represented as an adjacent point in the cortex. Some variability exists among cats in the extent and retinotopic representation of the visual field in area 17.

The distribution of substance P immunoreactive fibers in the rat central nervous system

Abstract: A detailed account of the distribution of immunoreactive substance P-containing structures in the rat central nervous system is presented, from results obtained by applying an indirect immunofluorescent technique. High densities of substance P-containing nerve terminals were present in sensory nuclei and other non-sensory structures such as thalamus, hypothalamus and extrapyramidal system. Substance P-reactive neuron cell bodies were present in spinal root ganglia, nucleus habenulae medialis, nucleus interpeduncularis, caudoputamen and globus pallidus. Most of the neocortex and the cerebellar cortices had no substance P-positive elements. The results support the hypothesis that substance P may be a widespread neurotransmitter in the central nervous system.

The afferent connections of the main and the accessory olfactory bulb formations in the rat: an experimental HRP-study.

Abstract: The afferent connections of the main and accessory olfactory bulbs in the rat were examined by injecting horseradish peroxidase (HRP) into one or the other of these structures either by microelectrophoresis or by hydraulic pressure. Alternate sections were stained with newly developed HRP-procedures using either benzidine dihydrochloride (de Olmos and Heimer, '77) or tetramethyl-benzidine. Eighteen to twenty-four hours after unilateral HRP injections confined to the main olfactory bulb, a large number of HRP-labeled perikaria appeared in the following telencephalic structures on the ipsilateral side: All portions of the anterior olfactory nucleus (AON) except its external part, the lateral transitional field (LT) between AON and the paleocortex, the whole extent of the primary olfactory cortex (POC); The medial forebrain bundle area deep to the olfactory tubercle, the nucleus of the horizontal limb of the diagonal band (NHDB) and the nucleus of the lateral olfactory tract (NLOT). A moderate to small number of labeled cells, furthermore, were seen in the dorsal (DT) and medial (MT) transition fields, the ventral praecommissural hippocampus (tt2), the ventral superficial part of the nucleus of the vertical limb of the diagonal band (NVDB), the sublenticular part of the substantia innominata (SI), the anterior amygdaloid area, the posterolateral cortical amygdaloid nucleus (C2) and the transition region (28 L') between the olfactory cortex and the lateral entorhinal area proper. On the contralateral side a large number of labeled cells were found in all parts of the AON, with especially heavy labeling in its external part. A moderate number of labeled cells could also be detected in the lateral transition field (LT) and the NLOT. In the diencephalon and the brain stem a moderate number of HRP-labeled perikaria were observed in the dorsal, perifornical, and lateral hypothalamus, as well as in locus coeruleus and the dorsal and medial raphae nuclei. Following large HRP injections in the main olfactory bulb a moderate to small number of labeled cells were seen also in the posterior and premammillary hypothalamus and in field CA1 of the retrocommissural hippocampus on the ipsilateral side, as well as in POC on the contralateral side. It is possible, however, that the uptake of label took place in an undetected pool of HRP in the very rostral part of AON rather than in the olfactory bulb. HRP injections in the accessory olfactory bulb resulted in labeled neurons in the posterior ventro-lateral part of the bed nucleus of the stria terminalis, the nucleus of the accessory olfactory tract, the rostrodorsal portions of the medial amygdaloid nucleus, and whole extent of the posteromedial cortical amygdaloid nucleus (C3) on the ipsilateral side. A few lightly labeled cells were seen also in the contralateral C3.

A description of the amygdaloid complex in the rat and cat with observations on intra‐amygdaloid axonal connections

Abstract: The cytoarchitectonic structure and divisions of the amygdaloid complex are described in the rat and cat, with special reference to the axonal connections of each of the amygdaloid nuclei, and to interspecies variations and similarities. Several intra-amygdaloid connections are also described, based on autoradiographic experiments with small injections of 3H-amino acids into the individual nuclei. Although it has probably not been possible to determine all of the intra-amygdaloid projections from these experiments, the connections which have been shown are very specific. The lateral nucleus projects to the basomedial nucleus, the lateral part of the central nucleus, and the periamygdaloid cortex, while the basolateral nucleus projects only to itself, the medial part of the central nucleus, and the nucleus of the lateral olfactory tract. The basomedial nucleus projects to the cellular layer of the medial nucleus and the amygdalo-hippocampal area, while the molecular layer of these structures and of the posterior cortical nucleus, receives projections from the periamygdaloid cortex or from the endopiriform cortex. There are also interconnections between the medial and posterior cortical nuclei, and commissural connections between the posterior cortical nuclei of the two sides.

An autoradiographic study of the organization of intrahippocampal association pathways in the rat.

Abstract: The longer associational connections of the hippocampal formation have been studied autoradiographically in a series of adult rats after small injections of 3H-amino acids into each of its various cytoarchitectonic fields. The major findings can be summarized as follows. The dentate gyrus projects in a topographically ordered manner upon the pyramidal cells of the regio inferior by way of the supra- and infrapyramidal bundles of mossy fibers. Certain cells in the hilar region of the dentate gyrus (which operationally may be defined as constituting field CA4 of Ammon's horn) give rise to a hippocampodentate projection to the inner one-quarter of the molecular layer of the dentate gyrus. Either the same or closely related cells give rise to fibers which join the Schaffer collateral system from field CA3 to the stratum radiatum and stratum oriens of the regio superior. The regio inferior is also characterized by a longitudinally directed associational bundle which runs throughout the septo-temporal extent of the hippocampus and is centered in the region of subfield CA3a. The regio superior has no reciprocal projection to the regio inferior but sends a substantial projection back to the subiculum and to the entorhinal area. There is also a projection to the subiculum from the regio inferior, and the subiculum itself probably contributes significantly to the projection to the entorhinal and perirhinal cortices. There is a striking parallelism between certain of these associational connections and the commissural projections to the hippocampus and dentate gyrus. Each cytoarchitectonic field that contributes a commissural projection also gives rise to an ipsilateral associational pathway which in its intrahippocampal course and its mode of termination exactly matches that of the commissural projection, although in general, the associational connections are more extensive in their distribution along the septo-temporal extent of the hippocampus than the corresponding commissural connections. The reverse is not true; there are a number of associational projections which are not paralleled by commissural projection. All of the associational projections are topographically arranged, but those which extend across the transverse axis of the hippocampus usually show considerable divergence so that afferents from different levels overlap fairly considerably within their respective projection fields.

Nuclei of the solitary tract: efferent projections to the lower brain stem and spinal cord of the cat.

Abstract: The efferent projections from the solitary complex to the lower brain stem and spinal cord were studied in the cat with the autoradiographic anterograde axonal transport and retrograde horseradish peroxidase (HRP) techniques. A revised cytoarchitectonic description of the caudal two-thirds of the complex is presented in which the complex was subdivided into six nuclei: lateral, ventrolateral, intermediate, medial, parvocellular, and commissural solitary tract nuclei. Following injections of 3H amino acids into electrophysiologically defined regions of the complex in which cardiac or respiratory units were recorded, labeled fibers could be traced to a number of sites in the caudal brain stem including the medial and lateral parabrachial nuclei, Kolliker-Fuse nucleus and the area ventral to this nucleus, lateral periaqueductal gray matter, ambiguus complex, which consists of the retrofacial, ambiguus and retroambiguus nuclei, ventrolateral reticular nucleus (in an area equivalent to the A1 cell group of Dahlstrom and Fuxe, '64), medial accessory olive, paramedian reticular formation, and lateral cuneate nucleus. Descending solitario-spinal projections have been traced bilaterally, but predominantly to the contralateral side, to the region of the phrenic motor neurons in the C4-C6 ventral horn, to the thoracic ventral horn, and intermediolateral cell column. Confirmatory evidence of some of these projections was obtained from a series of HRP experiments. Mainly small neurons of the parvocellular, medial and commissural solitary tract nuclei project to the region of the parabrachial and Kolliker-Fuse nuclei. The lateral solitary nucleus projects almost exclusively to the ipsilateral medial accessory olive. It was not possible to interpret conclusively the labeling seen in the medium and large neurons of the ventrolateral solitary nucleus after HRP injections made in the region of the ambiguus-retroambiguus complex due to the problem of fibers of passage. Following injections of HRP into the cervical, thoracic, lumbar, or sacral spinal cord, retrograde cell labeling was seen in the solitary complex. Cells in the intermediate and commissural nuclei were labeled after all four types of experiments. In the ventrolateral nucleus, medium sized neurons were predominantly labeled after the cervical spinal cord experiments, while large sized neurons were labeled mainly after the thoracic spinal cord injections. The potential physiological significance of these connections is discussed in terms of central control of cardiovascular and respiratory functions.

Association and commissural fiber systems of the olfactory cortex of the rat. I. Systems originating in the piriform cortex and adjacent areas

Abstract: The association and commissural fiber systems arising in the olfactory cortical areas caudal to the olfactory peduncle (the piriform cortex, nucleus of the lateral olfactory tract, anterior cortical nucleus of the amygdala, periamygdaloid cortex and entorhinal cortex) have been studied utilizing horseradish peroxidase as both an anterograde and a retrograde axonal tracer. In the piriform cortex two sublaminae within layer II (IIa and IIb) and layer III have been found to give rise to distinctly different projections. Retrograde cell labeling experiments indicate that the association fiber projection from layer IIb is predominantly caudally directed, while the projection from layer III is predominantly rostrally directed. Cells in layer IIa project heavily to areas both caudal and rostral to the piriform cortex. The commissural fibers from the piriform cortex are largely restricted in their origin to layer IIb of the anterior part of the piriform cortex and in their termination on the contralateral side to the posterior part of the piriform cortex and adjacent olfactory cortical areas. A projection to the olfactory bulb has also been found to arise from cells in layers IIb and III of the ipsilateral piriform cortex, but not in layer IIa. In addition to those from the piriform cortex, association projections have also been found from other olfactory cortical areas. The nucleus of the lateral olfactory tract has a heavy bilateral projection to the medial part of the anterior piriform cortex and the lateral part of the olfactory tubercle (as well as a lighter projection to the olfactory bulb); both the anterior cortical nucleus of the amygdala and the periamygdaloid cortex project ipsilaterally to several olfactory cortical areas. The entorhinal cortex has been found to project to the medial parts of the olfactory tubercle and the olfactory peduncle. The olfactory tubercle is the only olfactory cortical area from which no association fiber systems (instrinsic or extrinsic) have been found to originate. A broad topographic organization exists in the distribution of the fibers from several of the olfactory areas. This is most obvious in the anterior part of the olfactory cortex, in which fibers from the more rostral areas (the anterior olfactory nucleus and the anterior piriform cortex) terminate in regions near the lateral olfactory tract, while those from more caudal areas (the posterior piriform cortex and the entorhinal cortex) terminate in areas further removed, both laterally and medially, from the tract. Projections to olfactory areas from the hypothalamus, thalamus, diagonal band, and biogenic amine cell groups have been briefly described.

Three bulbospinal pathways from the rostral medulla of the cat: an autoradiographic study of pain modulating systems.

Abstract: Small amounts of 3H-leucine were injected into discrete regions in the rostral medulla of the cat. Descending projections from these sites were studied with autoradiographic methods. On the basis of differential projections to the medulla and spinal cord, three distinct regions were delineated. Nucleus reticularis gigantocellularis (Rgc), located dorsally in the medullary reticular formation, projects primarily to “motor” related sites, including cranial motor nuclei VI, VII, XII, nucleus intercalatus, and a part of the ipsilateral medial accessory olive. The projection to the spinal cord is primarily via the ipsilateral ventrolateral and contralateral ventral funiculi. The Rgc terminal field is in lamina VII and VIII ipsilateral and lamina VIII contralateral to the injection site. In contrast, nucleus raphe magnus, (NRM) located ventrally, in the midline of the rostral medulla projects primarily to structures with known nociceptive and/or visceral afferent input. These sites include the solitary nucleus, the dorsal motor nucleus (X) and the marginal and gelatinous layers of the spinal trigeminal nucleus caudalis. The projection to the spinal cord is bilateral, via the dorsolateral funiculus. Terminal fields are found in the marginal zone and the substantia gelatinosa of the dorsal horn, and more deeply in lamina V, medial VI and VII. Nucleus reticularis magnocellularis (Rmc), located lateral to NRM and ventral to Rgc, has an overlapping projection with NRM, but the projection is ipsilateral. This difference between Rmc and Rgc is correlated with cytoarchitectural features of the two regions. The possibility that the raphe-spinal pathway in the DLF mediates opiate and brain stimulation-produced analgesia is discussed.

Ocular dominance columns and their development in layer IV of the cat's visual cortex: a quantitative study.

Abstract: The distribution of geniculocortical afferents serving the left and right eyes was studied in abult cats and in kittens of various ages. Methods used were autoradiography of transneuronally transported 3H-proline injected into one eye, and physiological recordings. In the adult cat, patches of label in layer IV corresponding to ocular dominance columns were seen both ipsilateral and contralateral to the injected eye. Between the patches, however, grain density was substantially above background, especially on the contralateral side. Similarly, in the contralateral lateral geniculate nucleus (LGN) there was substantial labelling of neuronal cell bodies in lamina A1, which receives no innervation from the injected eye. Such spillover of radioactivity into the inappropriate laminae of the LGN was measured in autoradiographs of semithin sections, and its effect on the cortical labelling pattern calculated. Spillover appeared to account quantitatively for the labelling seen between patches in the cortex. It was concluded that the geniculocortical afferents for the ipsilateral and contralateral eyes are equally and almost completely segregated from each other at the centers of columns, although there is extensive overlap at the borders. This pattern is consistent with the physiological pattern of ocular dominance in layer IV. In kittens studied at one to two weeks of age, radioactive label formed a continuous band in layer IV on both sides of the brain. On the ipsilateral side, this appearance could not be accounted for by spillover in the LGN, but reflected a continuous, non-columnar distribution of afferents. Physiological recordings at this age showed most cells in layer IV to be nearly equally responsive to stimulation of either eye, in contrast to the adult pattern. Periodic variations in grain density were first noted at three weeks, and a pattern similar to that of the adult was reached by about six weeks of age. On the contralateral side the uniform labelling pattern seen in the 1- to 2-week-old kittens was uninterpretable, owing to the very great spillover of radioactivity in the contralateral LGN, but the physiology suggested that the contralateral afferents, too, were uniformly distributed in layer IV. The results suggest that the earliest functional connections formed by geniculocortical afferents have a uniform, non-columnar arrangement in layer IV, and that the formation of the adult pattern is likely to involve the breakage and reformation of synaptic connections. This process appears to be similar to that described for the monkey (Hubel et al., '77; Rakic, '76), except that the beginning of segregation is postponed until postnatal life.

The retinotopic organization of lateral suprasylvian visual areas in the cat.

Abstract: This is the second in a series of papers in which we describe our continuing efforts to define functional units of visual cortex based upon electro-physiological mapping of single and multiple unit activity in both awake and the nitrous oxide anesthetized cats. In the first paper (Tusa, Palmer and Rosenquist, '78), the extent and retinotopic organization of area 17 were described. In this paper, we describe the somewhat more complex organization of the visual cortex lying on the banks of the middle and posterior suprasylvian sulci. This region of cortex consists of six retinotopically organized units. These areas are arranged as three roughly mirror symmetrical pairs separated in each case by the fundus of the middle or posterior suprasylvian sulci. Some thalamo-cortical autoradiographic material is presented which supports this parcellation of the cortex.

Afferent projections to the rat locus coeruleus as determined by a retrograde tracing technique.

Abstract: Afferent connections to the rat locus coeruleus (LC), which contains exclusively noradrenergic neurons, have been traced using the technique of retrograde transport of horseradish peroxidase (HRP). In order to ensure accurate placement of adequate amounts of HRP in the LC, a microiontophoretic delivery technique coupled with single cell recording was employed. The use of electro physiological "landmarks" as aids in placing the injections is described. Following HRP injections into the LC, forebrain structures containing labelled neurons included the insular cortex, the central nucleus of the amygdala, the medial, lateral and magnocellular preoptic areas, the bed nucleus of the stria terminalis, and the dorsomedial, paraventricular and lateral hypothalamic areas. In the brainstem reactive neurons were observed in the central grey substance, the reticular formation, the raphe, vestibular, solitary tract and lateral reticular nuclei. In particular, the areas of catecholamine cell groups A1, A2 and A5 appeared to contain many reactive cells. Labelled neurons were also observed in the fastigial nuclei and in the marginal zones of the dorsal horns of the spinal cord. This pattern of afferent innervation supports suggestions for a role for the LC in behavioral arousal mechanisms and autonomic regulation.

Catecholamine innervation of the basal forebrain II. Amygdala, suprarhinal cortex and entorhinal cortex

Abstract: The catecholamine (CA) innervation of the posterior basal forebrain, the amygdala, suprarhinal cortex and entorhinal cortex, was studied in the rat using biochemical assay and fluorescence histochemistry. The assay studies demonstrate a moderate norepinephrine (NE) content in the amygdala and entorhinal cortex with a lower value for the suprarhinal cortex. Following destruction of the locus coeruleus, the decrease in NE content of these basal forebrain structures indicates that their principal NE innervation is from locus coeruleus. An additional small NE input arises from the medullary NE neuron groups. Ablation of dopamine (DA) cell groups (substantia nigra-ventral tegmental area, SN-VTA) indicates that the DA input to the amygdala arises from the lateral VTA and medial half of the SN. Fluorescence histochemical studies using the glyoxylic acid-Vibratome technique demonstrate the presence of four distinct types of CA neuron terminal plexus in the posterior basal forebrain. These include two different DA fiber types arising in SN-VTA, small NE fibers with small varicosities arising in locus coeruleus and NE fibers with larger varicosities arising in other brainstem NE cell groups. The large NE fibers appear to enter the amygdala via the ansa peduncularis-ventral amygdaloid bundle to innervate the central and basolateral nucleus and the anterior amygdaloid area. The locus coeruleus NE fibers appear to enter the posterior basal forebrain via both the stria terminalis and ansa peduncularis-ventral amygdaloid bundle system to form a moderately dense innervation of the central and basolateral nuclei of the amygdala and a less dense innervation of the other areas. The DA neuron axons are concentrated in the central and basal nuclei and intercalated cell groups. Other areas receive a more diffuse DA input, with the exception of the moderately dense innervation of the suprarhinal cortex and DA “islands” in the ventral-anterior entorhinal cortex. The DA input to the posterior basal forebrain is complex and heterogeneous and the axonal morphology differs greatly among the terminal fields within the amygdala and adjacent cortical areas.

Anatomical demonstration of orientation columns in macaque monkey

Abstract: In the macaque monkey striate (primary visual) cortex, the grouping of cells into ocular dominance and orientation columns leads to the prediction of highly specific spatial patterns of cellular activity in response to stimulation by lines through one or both eyes In the pesent paper these paterns have been examined by the 2-deoxyuglucose autoradiographic method developed by Sokoloff and his group (Kennedy et al, '76) An anesthetized monkey was given an injection of 14C 2-deoxyglucose and then visually stimulated for 45 minues with a large array of moving vertical stripes, with both eyes open The 14C autoradiographs of striate cortex showed vertical bands of label extending through the full cortical thickness Layer I was at most only lightly labelled, and layers IV b and VI wee the most dense Layer IV c (the site of terminations of most geniculate afferents) was labelled uniformly along its length, as expected from the lack of orientation specificity of units recorded in that leyer In the other layers the pattern seen in tangential sections was complex, consisting of swirling stripes with many bifurcations and blind endings, but with occasional more regular regions whee the stripes wee roughly parallel Interstripe distance was rather constant, at 570 μm Ocular dominance columns were examined in this same monkey, in the same region, by injecting one eye with 3H-proline two weeks before the deoxyglucose experiment, and preparing a second set of autoradiographs of the sections after prolonged washing to remove the 14C-deoxyglucose As seen in tangential sections through layer IV c, these columns had the usual stripe-like form, with a period of 770 μm, but were simpler in their pattern than the orientation stripes, with fewer bifurcations and less swirling A comparison of the two sets of columns in the same area showed many intersections, but no strict or consistent relationships: angles of intersection showed a distribution that was not obviously different from that expected for any two randomly superimposed sets of lines Another monkey was stimulated with vertical stripes, but with only one eye open Deoxyglucose autoradiographs of tangential sections showed regular uniform rows of label in layer IV c, with all the characteristic features of eye dominance columns In the layers above and below IV c the rows in tangential view were broken up into regularly spaced patches of label, presumably representing aggregations of cells responsive to vertically oriented stimuli The patches showed no consistent alignment across the ocular dominance rows, and indeed no such tendency would be expected, considering the complexity of the orientation columns This pattern of labelling is again predicted from and confirms the previous physiological studies

In situ injection of kainic acid: A new method for selectively lesioning neuronal cell bodies while sparing axons of passage

Abstract: The morphologic sequelae following stereotaxic injection into the rat striatum of kainic acid, a conformationally restricted analogue of glutamate, were examined by means of bright fields, histofluorescence and electron microscopic techniques. The neuropathologic response to kainate injection occurs in two distinct phases. Fist, the intrinsic neurons of the striatum undergo a rapid degeneration during the first 48 hours after injection; this is characterized by the sequential loss of cytoplasmic Nissl-substance (chromatolysis), shrinkage of the perikarya, clumping of the nuclear chromatin and finally disruption of the nuclear membrane. Between one and three weeks after injection, a marked proliferation of astrocytes in the gray matter formerly populated by neurons characterizes the second phase. The region of several neuronal loss in the kainate injected stratum is approximately spherical in shape, and its extent is non-linearly related to the amount of kainate injected. The neuropil of the injected striatum is markedly disrupted due to the death of intrinsic neurons and loss of their processes. Yet, histofluoresence microscopy demonstrates that the dopaminergic axons projecting from the substantia nigra do not degenerate in the kainate injected striatum; electron microscopic studies indicate that corticofugal fibers traversing the striatum also unaffected by kainate. Many presynaptic boutons, presumably of extrinsic origin, are intact up to ten days after injection; osmophilic vestiges of postsynaptic elements remain adherent to these boutons. Numerous phagocytic astrocytes are observed throughout the lesioned area. These morphologic studies provide (direct) evidence that in situ injection of kainic acid in brain causes a selective degeneration of neurons with cell bodies in the area of the injection but spares axons that arise from perikarya outside the region but pass through or terminate in the injected area. Thus, in situ injection of kainic acid is a new technique for making selective brain lesions that will be useful for examining neuronal connectivity.

Developmental studies of thalamocortical and commissural connections in the rat somatic sensory cortex

Abstract: Autoradiographic, axonal degeneration, and horseradish peroxidase fiber tracing methods were employed to investigate the organization, development and potential plasticity of the thalamocortical projection to the somatic sensory cortex of the rat. In the adult animal, thalamocortical terminals are concentrated primarily in layers I and IV and in the upper part of layer VI. Fibers terminating in layers IV and VI arise from a different thalamic region than those terminating in layer I. Discrete clusters of fibers and terminals 250–450 μm wide are distributed only to the parts of the SI cortex containing dense aggregates of layer IV granule cells and not to the intervening, less granular and commissurally connected zones. At birth, thalamocortical fibers have invaded the deep part of the developing SI cortex and are concentrated in the upper part of layer VI. Between the age of two and three days, an additional concentration of fibers appears in the part of the cortical plate which will become layer IV. Layer IV is clearly recognizable by three days of age and the dense granule cell aggregates appear in it no more than one day later. The ingrowth of commissural fibers (Wise and Jones, '76) lags behind that of thalamic fibers. The mature commissural fiber pattern is not established until the age of seven days. After removal of the developing thalamocortical system by thalamotomy in newborn rats, subsequent investigation of the commissural system in the adult showed that no commissural fibers or terminals had invaded either laminae or zones of the cortex deprived of thalamic input. Similarly, commissurotomy at birth was not followed by sprouting of thalamic fibers into zones or laminae deprived of commissural connections. The connectional specificity observed in these neocortical fiber systems contrasts markedly with the plasticity of connections reported in allocortical systems. Removal of thalamocortical afferents before they attain their definitive distribution does not radically effect the overall development of the dense granule cell aggregates in layer IV. Within the aggregates, however, subsidiary features such as the “barrels” fail to appear. This finding suggests that certain elements of cortical architecture such as the dense granule cell aggregates are independent of thalamic afferents while others, such as the barrels, result from the interaction of the developing thalamocortical fibers and/or terminals with maturing neurons.

Double representation of the body surface within cytoarchitectonic area 3b and 1 in “SI” in the owl monkey (aotus trivirgatus)

Abstract: Microelectrode multiunit mapping studies of parietal cortex in owl monkeys indicate that the classical "primary" somatosensory region (or "SI") including the separate architectonic fields 3a, 3b, 1, and 2 contains as many as four separate representations of the body rather than one, An analysis of receptive field locations for extensive arrays of closely placed recording sites in parietal cortex which were later related to cortical architecture led to the following conclusions: (1) There are two large systematic representations of the body surface within "SI." Each is activated by low threshold cutaneous stimuli; one representation is coextensive with Area 3b and the other with Area 1. (2) While each of these representations contains regions of cortex with topological or "somatotopic" transformations of skin surface, the representations have many discontinuities where adjoining skin surfaces are not adjoining in the representations. Thus, the representations can be considered as composites of somatotopically organized regions, but cannot be accurately depicted by simple continuous homunculi. Lines of discontinuity often cut across dermatomes and seldom follow dermatomal boundaries, i.e., neither cutaneous representation constitutes a systematic representation of dermatomal skin fields. (3) While the two cutaneous fields are basically similar in organization and are approxi- mate mirror images of each other, they differ in important details, i.e., lines of discontinuity in the representations and the sites of representations of dif- ferent specific skin surfaces differ significantly in the two representations. (4) The two cutaneous representations also differ in size and in the relative propor- tions of cortex devoted to representation of various body parts. Because the pro- portions in each representation differ, they cannot both be simple reflections of overall peripheral innervation density. (5) All or part of Area 2 contains a sys- tematic representation of deep body structures. These conclusions are consistent with a view of the anterior parietal region as containing functionally distinct fields at least partially related to different subsets of receptor populations and coding or representing different aspects of somatic sensation. We suggest that the "SI" region of primates be redefined as a parietal somatosensory strip, the Area 1 representation as the posterior cuta- neous field, and, for reasons of probable homology with "SI" of other mammals, the Area 3b representation as SI proper.

The connections of the nucleus reuniens thalami: Evidence for a direct thalamo-hippocampal pathway in the rat†

Abstract: The afferent and efferent connections of the rat's midline nucleus reunions thalami (reuniens) were studied by experiments using the methods of retrograde cell marking by horseradish peroxidase (HRP) and anterograde fiber tracing by autoradiography. A microelectrophoretic deposit of tritiated amino acids in reuniens provided the first evidence of a direct thalamo-hippocampal connection. Labeled reuniens efferents ascend to the genu of the corpus callosum and turn caudally in the cingulate fasciculus, from which fibers distribute to layer I of the anterior medial, cingulate, and retrosplenial cortices. A longer component of this system curves around the callosal splenium and forms a massive rostrally directed fiber sheet that innervates entorhinal and parahippocampal areas and Ammon's horn. Entorhinal afferents are localized to layers I and III, whereas the hippocampal afferent plexus is remarkably restricted to the stratum lacunosum-moleculare of the CA1 field and the corresponding stratum of the ventral subiculum. Reuniens projects more sparsely and diffusely to many subcortical structures, a number of which lie in the limbic domain: the anterior olfactory nucleus, nucleus accumbens, olfactory tubercle, amygdala, claustrum, septum, preoptic area, medial and lateral hypothalamic regions, deep portions of the pretectum and superior colliculus, rostral levels of the ventral tegmental area and central gray substance and, perhaps, the median eminence. The efferent connections of reuniens were examined with HRP. HRP deposited in the nucleus labeled small to moderate numbers of neurons in many structures extending from the frontal cortex to caudal midbrain levels. The appearance of cell labeling in regions projected upon by reuniens suggests a reciprocity of connections between it and the medial cortex, septum, preoptic area, amygdala, medial and lateral hypothalamic regions, ventral tegmental area, central gray substance, pretectum, superior colliculus and the subiculum. Cell labeling in regions not receiving its efferents – the ventral thalamus, midbrain tegmentum, mesencephalic raphe, and parabrachial nuclei – may hold another clue to the future understanding of the role of the nucleus reuniens in limbic functions.

The neurological organization of pathways between the dorsal lateral geniculate nucleus and visual cortex in old world and new world primates

Abstract: Pathways between the dorsal lateral geniculate nucleus (dLGN) and visual cortex in Old World (Macaca, Papio, Erythrocebus, Cercopithecus) and New World (Saimiri, Cebus) primates were studied after injections of horseradish peroxidase and H3 or S35 amino acids into the dLGN or visual cortex. Trans-synaptic autoradiography was also used to study these pathways after an injection of H3 proline-fucose into one eye. The subsequent autoradiographs of visual cortex showed that Old World primates have separate eye inputs (ocular dominance columns) in striate cortex, whereas New World monkeys have overlapping or non-separated eye inputs. In both primate groups the geniculocortical input to layer IVA formed a pattern which resembled a honeycomb in tangential sections, unlike the solidly labeled layer IVC. Also common to the two primate groups was a projection from dLGN to layer VI. There was no dLGN projection to any prestriate area in any of the primates. However, after an injection limited to the prestriate cortex of Macaca, light autoradiographic labeling was seen in the interlaminar zones and the magnocellular and S laminae, demonstrating a prestriate-dLGN pathway. Our results indicate that the primate visual system differs significantly from the cat in having no dLGN projection to area 18. There are also significant differences between primates in the level at which the possibility of binocularity (of an excitatory nature) first occurs in the striate cortex because in the species studied thus far with neuroanatomical methods, Old World primates have ocular dominance columns in layer IV but most New World monkeys lack them.

Prenatal development of the cerebellar system in the rat. I. Cytogenesis and histogenesis of the deep nuclei and the cortex of the cerebellum

Abstract: Prenatal cerebellar development was investigated with three approaches. In normal embryos sectioned in three planes morphological and cytological changes were determined at daily intervals beginning on embryonic day 13 (E13). A similar series of X-irradiated embryos was used to study changes in neuroepithelial organization and in the location of primitive (radiosensitive) or differentiated cells. Finally, to quantify the time of origin of different classes of cerebellar neurons with the progressively delayed labelling procedure, we used autoradiograms from adult rats whose mothers were injected with two successive daily doses of 3H-thymidine on overlapping days from day E13 on. The cerebellar anlage was delineated in the dorsal metencephalon by the collapse of its ventricular lining after X-irradiation. This “collapsing neuroepithelium” was located laterally on day E13, then it spread medially and reached the midline on day E16. Deep nuclear neurons began to differentiate on day E13, with two-thirds forming on day E14; Purkinje cell formation peaked on day E15, with a few cells still forming on day E16. It was postulated that the deep nuclear neurons settled first in the superficial “nuclear zone,” and that the Purkinje cells gathered temporarily in the underlying “transitory zone,” adjacent to the collapsing neuroepithelium. In the next period of cerebellar development four major events were recognized. (1) Beginning on day E17 the cells of the nuclear and transitory zones became intermingled. It was postulated that the Purkinje cells were migrating radially through the ranks of the stationary deep nuclear neurons and assembled under the spreading canopy of a fibrous plexus and the external germinal layer. (2) It was also on day E17 that the external germinal layer began to form as one of the prongs of the “germinal trigone” in the posteroventral aspect of the cerebellum. On the succeeding days the external germinal layer spread over the surface of the cerebellum; in the vermis in a rostral direction. (3) Two cell types destined to settle in the future granular layer, the pale cells and the Golgi cells, began to form at a relatively slow rate on day E19. Chronological considerations suggested that they were generated in the regressing, noncollapsing neuroepithelium of the cerebellar ventricle. (4) From the beginning (day E17) of its genesis posteroventrally, the primitive cerebellar cortex bridged the midline. As the fused cortex spread rostrally, the vertical ventricular cleft separating the underlying portions of the cerebellum became shallower and then disappeared; the process was completed in the anterior cerebellum by day E22. By the time of birth the maturation of the neurons of the deep nuclei appeared advanced but the maturation of the prenatally produced neurons of the cortex does not start until after birth when a new class of neurons is generated in the external germinal layer.

Association and commissural fiber systems of the olfactory cortex of the rat II. Systems originating in the olfactory peduncle

Abstract: The structure and connections of areas within the olfactory peduncle (anterior olfactory nucleus and tenia tecta) have been examined. The anterior olfactory nucleus has been divided into external, lateral, dorsal, medial, and ventro-posterior parts. In spite of the term nucleus which is applied to these areas, all of them contain pyramidal-type cells with apical and basal dendrites oriented normal to the surface, and are essentially cortical in organization. Experiments utilizing retrograde and anterograde axonal transport of horseradish peroxidase (HRP) have demonstrated that each of these parts of the anterior olfactory nucleus possesses a unique pattern of afferent and efferent connections with other olfactory areas. All subdivisions have projections to both the ipsilateral and contralateral sides, although the ipsilateral projection of the pars externa (to the olfactory bulb) is extremely light. Interestingly, crossed projections are in each case directed predominantly to areas adjacent to the homotopic areas. Two primary subdivisions may also be distinguished in the tenia tecta: a dorsal part composed largely of tightly packed neurons which closely resemble the granule cells of the dentate gyrus (bushy apical but no basal dendrites) and a ventral part which contains predominantly pyramidal-type cells. The connections of these two parts are also very different. The ventral tenia tecta receives substantial projections from the olfactory bulb, pars lateralis of the anterior olfactory nucleus, piriform cortex and lateral entorhinal area. It gives off a heavy return projection to the pars lateralis and lighter projections to the olfactory bulb, piriform cortex and olfactory tubercle. The dorsal tenia tecta receives a heavy projection from the piriform cortex, but none from the olfactory bulb. A few cells in the dorsal tenia tecta are retrogradely labeled from HRP injections into the medial aspect of the olfactory peduncle (involving the ventral tenia tecta and adjacent areas), but none are labeled from the other olfactory areas that have been injected. An area on the dorsal aspect of the olfactory peduncle that differs significantly from the anterior olfactory nucleus, tenia tecta and piriform cortex in terms of its connections and cytoarchitecture has been termed the dorsal peduncular cortex. The most striking feature of this area is its very heavy reciprocal connection with the entorhinal cortex, although it is also reciprocally connected with the olfactory bulb and piriform cortex and projects to the olfactory tubercle. Cells in layer I of the medial and ventral aspects of the olfactory peduncle have been retrogradely labeled from HRP injections into the olfactory tubercle and lateral hypothalamic area. These cells overlie the ventral tenia tecta, medial part of the anterior piriform cortex and pars ventro-posterior and pars lateralis of the anterior olfactory nucleus, but do not appear to be distributed in relation to the cytoarchitectonic boundaries. Possible functional roles of the areas within the olfactory peduncle have been discussed.

GABA-ergic pathways in the goldfish retina.

Abstract: A high-affinity uptake mechanism for [3H]-gamma-aminobutyric acid (GABA) has been localized to type H1 cone horizontal cells and type Ab pyriform amacrine cells in the retina of the goldfish by light and electron microscopy autoradiography. By stimulating isolated retinas with colored lights during incubation we have been able to use [3H]-GABA uptake as a probe of light-evoked changes in membrane potential. All colors of lights increase and darkness decreases [3H]-GABA uptake by H1 cone horizontal cells. Our model of voltage dependence of GABA uptake predicts that all colors of light should hyperpolarize H1 cone horizontal cells and other investigators have shown by intracellular recording and dye-marking that type H1 cone horizontal cells hyperpolarize to all wavelengths of light. We have also obtained evidence that dark-induced depolarization of cone horizontal cells leads to release of GABA. Type Ab pyriform amacrine cells show maximal [3H]-GABA uptake in darkness and when exposed to green or blue lights, but red lights dramatically suppress uptake. We predict these neurons to be red-depolarizing, and recent intracellular recordings and dye-marking by Famiglietti et al. ('77) support our conclusions. Synaptic relations of apparently GABA-ergic neurons were investigated in the electron microscope. We propose type H1 cone horizontal cells to be both pre- and post-synaptic to red-sensitive cones and type Ab pyriform amacrine cells to be both pre- and post-synaptic to red-sensitive center-depolarizing bipolar cells.

The axonal arborizations of lateral geniculate neurons in the striate cortex of the cat

Abstract: Horseradish peroxidase (HRP) was injected into the optic radiations of adult cats. With placements close to the lateral geniculate nucleus (LGN), the enzyme diffused retrogradely along the axons of geniculocortical relay neurons, entered their cell bodies, and, after reaction with diaminobenzidine, produced a Golgi-like staining of entire neurons. When the injections were made close to the visual cortex, the enzyme diffused anterogradely and filled complete axonal arborizations in area 17. In the LGN, examples of type 1 and type 2 relay neurons (Guillery, '66) were reconstructed, and their axon diameters measured. The type 1 neurons (thought to correspond to Y-cells – LeVay and Ferster, '77) possessed large diameter axons (2–3.3 μm), while the type 2 neurons (thought to be X-cells) had medium-sized axons (1–1.7 μm). Both types of neurons gave off axon collaterals to the perigeniculate nucleus. In the cortex, two types of afferent supplied layer IV. One distributed to the upper part of the layer (layer IVab), extending a short distance into layer III. The parent trunks of these axons, measured in the white matter, had diameters matching those of type 1 LGN relay cells. The other type distributed to layer IVc. The diameters of these axons matched those of type 2 LGN relay cells. Most afferents of both types gave off collaterals to layer VI — there were no axons which exclusively innervated this layer. The axons supplying layer IVab had a wide lateral spread in the cortex (up to 2 mm), and the boutons were grouped into two to five clumps, whose size and arrangement were similar to ocular dominance columns. The axons supplying layer IVc had a much more restricted arborization, usually consisting of a single clump of boutons. LGN neurons with very fine axons (less than 1 μm) were found in laminae C1–C3. They probably corresponded to Guillery's type 4 neurons. In the cortex, fine-diameter axons arborized in the upper half of layer I. These axons sometimes had collaterals in the lower part of layer III and in layer V. Taken together, the arborizations of the cortical afferents observed in the present study account fully for the autoradiographic labelling pattern seen after 3H-proline injections into the LGN (LeVay and Gilbert, '76). The identification of type 1 and type 2 neurons as Y- and X-cells is strengthened by the observed difference in their axon diameters, in agreement with the different axonal conduction velocities reported for Y- and X-cells. The presence of cells with very fine axons in the deeper C laminae is consistent with the reported presence of W-cells (which have slowly conducting axons) in these layers. We conclude that the different classes of geniculate relay neuron have different laminar projections in area 17.

Golgi studies of the neurons in layer II of the dorsal horn of the medulla (trigeminal nucleus caudalis)

Abstract: Two kinds of pyramidal neuro ns and two kinds of multipolar neurons have been identified in layer I in the adult cat on the basis of thier dendritic morphology. The spiny pyramid emits an apical dendrite which beings dividing within 50 μm of the cell body and gives rise to an extensive dendritic arbor in which some second and third order branches extend for over 100 μm. The primary branches of the basal dendrites divide into secondary branches within 20 μm of the cell body. At each of these divisions, one daughter secondary branch will turn 180° and course in the direction of the apical dendrite. It then undergoes further branching and becomes intermingled in the apical dendritic arbor. These recurrent basal dendrites and the apical dendrites are charcterized by numerous long-necked spines. The other secondary basal branch either courses at right angles to the apical dendrite or droops basally away from it. Many of these basal dendrites are not as robust and extensive as the other dendritic branches. They may end without further branching or when they do branch they tend to be thinner and shorter with some showing signs of stunting. Smooth pyramids differ from the spiny pyramids in several respects. Their dendritic arbors contain fewer, more widely spaced spines and are more expansive than those of the spiny pyramids. While they still send branches into the apical arbors many branches run basally away from the cell body for considerable distances to give the entire dendritic field a more circular appearance. The compact multipolar neuron is found in the inner half of layer I. The long axis of its compact dendritic arbor (∼ 50 μm wide) is oriented in the rostrocaudal axis of the layer. Most of its dendritic branches lie to one side of the cell body squeezed between the axon bundles of the inner half of layer I. The loose multipolar neuron in found throughout the layer. Its loose dendritic arbor is at least 200 μm wide and, like the spiny pyramid, gives rise to several dendrites which terminate abruptly with little branching and without much change in diameter. The dendrites of all four layer I neurons are confined within layer I and its extensions into the spinal V tract. These four layer I neurons are considered to be Golgi type I projection neurons on the basis of the morphology of their initial axonal segment. Golgi type II inteneurons are not found in layer I.