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Projections of the nucleus of the tractus solitarius in the pigeon (Columba livia).

With the aid of autoradiographic and histochemical (WGA-HRP) tracing techniques, the projections of the nucleus of the tractus solitarius (nTS) in the pigeon have been delineated and related to the viscerotopic organization of the nucleus. As in mammals, nTS projects to both brainstem and forebrain structures. At medullary levels, projections were seen to nTS itself, to the dorsal motor nucleus of the vagus and to the subjacent and more ventral reticular formation. There is a substantial projection to the parabrachial nuclear complex with terminations in all its subnuclei and minor projections to locus coeruleus and several mesencephalic areas, including the ventral area of Tsai, the nucleus of the ascending brachium conjunctivum, and the compact portion of the tegmental pedunculopontine nucleus. At diencephalic levels, projections to the hypothalamus (magnocellular periventricular nucleus, stratum cellulare internum and externum) and dorsal thalamus were seen. Terminal fields within the basal telencephalon included the nucleus of the pallial commissure, the bed nucleus of the stria terminalis, and the nucleus accumbens. The organization of nTS projections in pigeons is correlated with the pattern of inputs to specific nTS subnuclei. Lateral tier subnuclei receiving cardiovascular and pulmonary inputs project upon the ventrolateral reticular formation and the ventrolateral parabrachial complex. Medial tier subnuclei receiving gustatory and gastrointestinal inputs project upon dorsal and medial parabrachial nuclei. Transparabrachial projections arise from nTS subnuclei receiving little or no primary input from the viscera.

Animals↗

Glycine-immunoreactive projection of the cat lateral superior olive: possible role in midbrain ear dominance.

Neurons in the lateral superior olive are optimally excited by stimulation of the ipsilateral ear, as are those in the inferior colliculus by stimulation of the contralateral ear. This reversal of ear dominance may result, in part, from distinct crossed excitatory and uncrossed inhibitory pathways ascending from the lateral superior olive. To explore this possibility, immunoreactivity for two putative inhibitory neurotransmitters, glycine and GABA, was examined in projection neurons that retrogradely transported horseradish peroxidase from the cat inferior colliculus. The results suggest that the projection from the lateral superior olive can be segregated, immunocytochemically, into three components: 1) a crossed, glycine-negative (-) projection; 2) an uncrossed, glycine-positive (+) projection; and 3) an uncrossed, glycine(-) projection. Additional evidence suggests that the terminal fields of the two uncrossed projections may distribute differently within the inferior colliculus. Glycine(+) or glycine(-) projection neurons, crossed or uncrossed, do not differ in the size, shape, or location of their somata. However, most glycine(-) neurons are heavily encrusted with glycine(+) endings; glycine(+) neurons have 40-60% fewer of these endings. Glycine(-) neurons located in the lateral limb have fewer glycine (+) perisomatic endings than those in the medial limb. Few projection neurons are GABA(+), and GABA(+) perisomatic endings are rare in the lateral superior olive. Thus, there is a heavy uncrossed projection from the cat lateral superior olive to the inferior colliculus that may be glycinergic and inhibitory. Furthermore, there is a bilateral projection that is not glycinergic or GABAergic, which may be excitatory. The potential contribution of these pathways to contralateral ear dominance in the inferior colliculus is discussed.

Animals↗

Topographical organization of the projections from the reticular thalamic nucleus to the intralaminar and medial thalamic nuclei in the cat.

The topography of the projections from the reticular nucleus of the thalamus (RT) to the intralaminar and medial thalamic nuclei were studied in the cat by the method of retrograde transport of horseradish peroxidase (HRP). Single small injections of the enzyme were made in the different intralaminar nuclei--mediodorsal, ventromedial, midline, and habenular--and in anterior group nuclei. The location and density of the neuronal labeling in the different parts of the RT were studied in each case. Our results show that 1) after injections located in all the nuclei here studied, a consistent number of labeled neurons were found in the RT, except for the injections in the lateral habenula and the anterior thalamic nuclear complex, both of which did not label neurons in the RT. 2) Among the other thalamic nuclei here studied, the most medially situated receive less numerous RT projections than those most laterally located. 3) Injections in all the nuclei studied gave rise to a cellular labeling in the anterior sectors of the RT, except for the anterior nuclear group and the lateral habenula. The projections from the rostral pole of the RT were topographically mediolaterally organized. 4) The anterodorsal part of the pregeniculate sector of the RT projects upon the large-celled part of the lateral central nucleus and to a lesser extent upon the paracentral, centromedian, and ventromedial nuclei, the anterior part of the lateral central nucleus, and the lateral band of the mediodorsal nucleus. The posterodorsal part of the RT pregeniculate sector only projects to the large-celled part of the lateral central nucleus. The dorsal portion of the posteroventral part of the RT pregeniculate sector also projects upon the large-celled part of the lateral central nucleus; its ventral portion projects to the ventromedial nucleus, the posterior part of the paracentral nucleus, the lateral band of the mediodorsal nucleus, and the centromedian nucleus. 5) The infrageniculate sector of the RT projects to the posterior part of the ventromedial nucleus. A weaker projection to the large-celled part of the lateral central nucleus, the centromedian nucleus, and the lateral band of the mediodorsal nucleus was also observed. 6) The ventral lateral geniculate nucleus projects upon the large-celled part of the lateral central nucleus, the lateral band of the mediodorsal nucleus, and the ventromedial nucleus. All these findings suggest an important modulatory action of the RT on the activity of the thalamic nuclei considered here.

Animals↗

Visual projections to the pontine nuclei in the rabbit: orthograde and retrograde tracing studies with WGA-HRP.

Visual projections to the pontine nuclei in the rabbit were examined by means of both orthograde and retrograde tracing of WGA-HRP. The tecto-pontine projection was examined following microinjections of WGA-HRP in the right superior colliculus. The projection to the pontine nuclei is strictly ipsilateral and terminates at middle and caudal levels of the pons. The projection is absent in rostral pontine nuclei. The strongest projection is to the dorsal border of the dorsolateral pontine nuclei and is the only projection seen when the primary injection site is confined to superficial laminae. When the primary injection site also includes intermediate and deep laminae, patches of labelled terminals are also seen within dorsolateral, lateral, peduncular, paramedian, and ventral pontine nuclei as well as in the contralateral nucleus reticularis tegmenti pontis. The striate corticopontine projection was also examined with orthograde tracing of WGA-HRP. The striate corticopontine projection is ipsilateral. Most labelled terminals were seen in dorsolateral and lateral pontine nuclei throughout the rostral half of pons with some additional terminal labelling in paramedian and peduncular nuclei. Labelled terminals were also seen in ventral pontine nuclei throughout the middle and caudal levels of the pons. In a retrograde tracing study, visual projections to the pontine nuclei were examined following microinjections of WGA-HRP into the pontine nuclei. Labelled cells were seen ipsilaterally in superficial and deep laminae of the superior colliculus and in layer V of striate and surrounding occipital cortex. The pontine nuclei also receive ipsilateral projections from the ventral lateral geniculate, the nucleus of the optic tract, anterior and posterior pretectal nuclei, and the dorsal and medial terminal nuclei of the accessory optic system. These pathways are potential sources of visual input to the cerebellum.

Animals↗

Cortical projection patterns of the medial septum-diagonal band complex.

A detailed analysis of the cortical projections of the medial septum-diagonal band (MS/DB) complex was carried out by means of anterograde transport of Phaseolus vulgaris leucoagglutinin (PHA-L). The tracer was injected iontophoretically into cell groups of the medial septum (MS) and the vertical and horizontal limbs of the diagonal band of Broca (VDB and HDB), and sections were processed immunohistochemically for the intra-axonally transported PHA-L. The labeled efferents showed remarkable differences in regional distribution in the cortical mantle dependent on the position of the injection site in the MS/DB complex, revealing a topographic organization of the MS/DB-cortical projection. In brief, the lateral and intermediate aspects of the HDB, also referred to as the magnocellular preoptic area, predominantly project to the olfactory nuclei and the lateral entorhinal cortex. The medial part of the HDB and adjacent caudal (angular) part of the VDB are characterized by widespread, abundant projections to medial mesolimbic, occipital, and lateral entorhinal cortices, olfactory bulb, and dorsal aspects of the subicular and hippocampal areas. Projections from the rostromedial part of the VDB and from the MS are preponderantly aimed at the entire hippocampal and retrohippocampal regions and to a lesser degree at the medial mesolimbic cortex. Furthermore, the MS projections are subject to a clear mediolateral topographic arrangement, such that the lateral MS predominantly projects to the ventral/temporal aspects of the subicular complex and hippocampus and to the medial portion of the entorhinal cortex, whereas more medially located cells in the MS innervate more septal/dorsal parts of the hippocampal and subicular areas and more lateral parts of the entorhinal cortex. PHA-L filled axons have been observed to course through a number of pathways, i.e., the fimbria-fornix system, supracallosal stria, olfactory peduncle, and lateral piriform route (the latter two mainly by the HDB and caudal VDB). Generally, labeled projections were distributed throughout all cortical layers, although clear patterns of lamination were present in several target areas. The richly branching fibers were abundantly provided with both "boutons en passant" and terminal boutons. Both distribution and morphology of the labeled basal forebrain efferents in the prefrontal, cingulate, and occipital cortices closely resemble the distribution and morphology of the cholinergic innervation as revealed by immunohistochemical demonstration of choline acetyltransferase. In contrast, the labeled projections to the olfactory, hippocampal, subicular, and entorhinal areas showed a heterogeneous morphology. Here, the distribution of only the thin varicose projections resembled the distribution of cholinergic fibers.

Animals↗

Development of retinotopy in projections from the eye to the dorsal lateral geniculate nucleus and superior colliculus of the wallaby (Macropus eugenii).

The development of retinotopy in projections from the eye to the dorsal lateral geniculate (dLGN) and superior colliculus (SC) has been studied in the marsupial wallaby. Discrete retinal lesions were made and the remaining retinal projections were traced with horseradish peroxidase in animals at stages ranging from just after optic innervation of the dLGN and SC to the time when the projections are mature. Topographically organised projections could be recognized a few weeks after axons first reached the dLGN and SC with a topographically discrete projection from nasoventral retinal recognized later than from dorsal, dorso-temporal, temporal, and temporoventral retina. Over time there was an increase in precision of the retinotopy as judged by an increase in sharpness of the borders of filling defects in the projection labelled with horseradish peroxidase. Refinement of the projection from temporal retina preceded that from nasal retina in both the dLGN and SC and in the former occurred concomitantly with the segregation of eye-specific terminal bands. Refinement was complete 16 weeks after birth, prior to eye opening at around 20 weeks after birth. Inequalities in retinal representations in both nuclei were present from the time retinotopy could first be detected. This was before the inequalities in retinal ganglion cell distribution, which underly these representations in the adult, were obvious. Retinotopy and inequalities in retinal representation characteristic of the adult are present from a very early stage in the protracted development of visual projections in the wallaby. Refinement may involve death of inappropriately projecting cells, pruning of inappropriately projecting axon arborizations or could be achieved by growth of the retinorecipient neuropil. Temporonasal differences in the time course of refinement may reflect gradients of maturation in the retina.

Animals↗

Organization of intrahippocampal projections originating from CA3 pyramidal cells in the rat.

The distribution of intrahippocampal projections arising from the CA3 region of the rat hippocampus was investigated using in vitro and in vivo methods. In the in vitro hippocampal slice preparation, single CA3 pyramidal cells were intracellularly labeled with horseradish peroxidase (HRP), and the three-dimensional organization of the axonal plexus was analyzed by using a computer-aided digitizing system. As many as eight primary collaterals originated from the principal axon of CA3 pyramidal cells and these commonly bifurcated further and innervated stratum oriens and stratum radiatum of CA3 and CA1. Within the 400 microns slice, the summed length of all visible collaterals per neuron ranged from 2.6 mm to approximately 12.5 mm. While the CA3 principal axon tended to be relatively smooth, the axonal collaterals bore numerous varicosities that electron microscopy confirmed to be presynaptic boutons. These varicosities occurred, on average, once every 7 microns of collateral length. The distribution of axonal collaterals differed depending on the location of the parent pyramidal cell. Only rarely could CA3 collaterals be followed in the slice to their terminations within CA1. To study the topographic organization of CA3 projections both to other levels of CA3 and to CA1, the anterograde tracer, Phaseolus vulgaris leucoagglutinin (PHA-L) was injected into various transverse and septotemporal levels of CA3. Immunohistochemical visualization of the lectin was conducted in dissected and "extended" hippocampi to facilitate analysis of the topographic distribution of projections along the long or septotemporal axis. Projections from all portions of CA3 reached widespread regions of CA3, CA2, and CA1, but only a few fibers entered the subicular complex and there were no projections to the entorhinal cortex. There were also some CA3 and CA2 projections to the hilus of the dentate gyrus, but these did not enter the granule cell or molecular layers. The CA3 projections to CA1 were organized according to several distinctive and consistent gradients that can generally be summarized as follows. 1. CA3 cells located close to the dentate gyrus (proximal CA3), while projecting both septally and temporally, tended to project more heavily to levels of CA1 located septal to the injection site. CA3 cells located closer to CA1, in contrast, projected more heavily to levels of CA1 located temporally to the injection site. 2. At, or close to, the septotemporal level of the injection, cells located proximally in CA3 gave rise to collaterals that tended to terminate more superficially in stratum radiatum than did those arising from mid and distal levels of CA3.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Organization of the thalamostriatal projections in the rat, with special emphasis on the ventral striatum.

The organization of the thalamic projections to the ventral striatum in the rat was studied by placing injections of the retrograde tracer cholera toxin subunit B in the ventral striatum and small deposits of the anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) in individual midline and intralaminar thalamic nuclei. In order to provide a complete map of the midline and intralaminar thalamostriatal projections, PHA-L injections were also made in those parts of the intralaminar nuclei that project to the dorsal striatum. The relationship of thalamic afferent fibres with the compartmental organization of the ventral striatum was assessed by combining PHA-L tracing and enkephalin immunohistochemistry. The various midline and intralaminar thalamic nuclei project to longitudinally oriented striatal sectors. The paraventricular thalamic nucleus sends most of its fibres to medial parts of the nucleus accumbens and the olfactory tubercle, whereas smaller contingents of fibres terminate in the lateral part of the nucleus accumbens and the most ventral, medial, and caudal parts of the caudate-putamen complex. The projections of the parataenial nucleus are directed towards central and ventral parts of the nucleus accumbens and intermediate mediolateral parts of the olfactory tubercle. The intermediodorsal nucleus projects to lateral parts of the nucleus accumbens and the olfactory tubercle and to ventral parts of the caudate-putamen. The projection of the rhomboid nucleus is restricted to the rostrolateral extreme of the striatum. A diffuse projection to the ventral striatum arises from neurons ventral and caudal to the nucleus reuniens rather than from cells inside the nucleus. Fibres from the central medial nucleus terminate centrally and dorsolaterally in the rostral part of the nucleus accumbens and medially in the caudate-putamen. Successively more lateral positions in the caudate-putamen are occupied by fibres from the paracentral and central lateral nuclei, respectively. The lateral part of the parafascicular nucleus projects to the most lateral part of the caudate-putamen, whereas projections from the medial part of this nucleus terminate in the medial part of the caudate-putamen and in the dorsolateral part of the nucleus accumbens. Furthermore, a rostral to caudal gradient in a midline or intralaminar nucleus corresponds to a dorsal to ventral and rostral to caudal gradient in the striatum. In the ventral striatum, thalamic afferent fibres in the "shell" region of the nucleus accumbens avoid areas of high cell density and weak enkephalin immunoreactivity.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Morphology and somatotopic organisation of the central projections of afferents from tactile hairs on the hind leg of the locust.

The morphology and organisation of the central projections of tactile hair afferents from the hind leg of the locust, Schistocerca gregaria, were examined by staining individual hair afferents. Each tactile hair on the femur, tibia, and tarsus is innervated by a single sensory afferent, which projects to the ipsilateral half of the metathoracic ganglion. Afferents arborize in the ventralmost and lateral ventral association centres (vVAC and lVAC). The projections are organised somatotopically in a map with three axes, according to the position of the hair on the leg. First, proximo-distal: afferents from hairs on the proximal leg segments project more anteriorly than do those from hairs on distal leg segments. Moreover, on any given segment the afferents from the more proximal hairs project more anteriorly than do the afferents from the distal hairs. Second, antero-posterior: afferents from hairs on the posterior surface of the leg project more medially than do afferents from anterior hairs. Third, dorso-ventral: afferents from hairs on ventral parts of the leg project more ventrally than do afferents from the dorsal hairs. The afferents from posterior and anterior hairs project to an area between the central projections from dorsal hairs and ventral hairs. The position of a projection within the map is dependent upon the location of the hair on the leg and not the peripheral routes taken by the axon of its afferent to reach the ganglion.

Animals↗

Lateral geniculate projections to the superficial layers of visual cortex in the tree shrew.

Our recent studies of tree shrew striate cortex have focused on the organization of lateral geniculate projections to layer IV and the projections from IV to layer III. Although these pathways play an important role in determining the response properties of layer III neurons, there are additional pathways from the lateral geniculate nucleus (LGN) that terminate directly in layer III. Previous studies provided evidence that these projections originate from layers 6 and 3 of the LGN and terminate in different subdivisions of layer III. In this study we used injections of biocytin to examine the projections of layers 6 and 3 to the cortex in more detail. Consistent with earlier work, we found that LGN layer 6 projects heavily to lower IIIc, while LGN layer 3 terminates densely in layer IIIb and sparsely throughout layers IIIa-I. In addition, we found that neurons in layers 6 and 3 have collateral projections: neurons in LGN layer 6 project to the bottom of layer IVb and sparsely to I-IIIb; neurons in LGN layer 3 project sparsely to layers V and VI and to the middle of IV. These patterns of projections are significant in the light of our studies of the connections from cortical layer IV to layer III. LGN projections to the superficial layers are organized into parallel pathways that exert selective influence over different populations of neurons in layers I-III and on the layer IV neurons that supply them.

Animals↗

Afferent projections to the mammillary complex of the rat, with special reference to those from surrounding hypothalamic regions.

To better understand the functional organization of the mammillary nuclei, we investigated the afferents to this nuclear complex in the rat with iontophoretically injected wheat germ agglutinin conjugated to horseradish peroxidase. Particular attention was paid to tracing local hypothalamic afferents to these nuclei. Injections into the medial mammillary nucleus (MMN) revealed strong projections from the subicular region, and weaker projections from the prefrontal cortex, medial septum, and the nucleus of the diagonal band of Broca. Other descending subcortical projections to the MMN arise from the anterior and the lateral hypothalamic area, the medial preoptic area, and the bed nucleus of the stria terminalis. Ascending afferents to the MMN were found to originate in the raphe and various tegmental nuclei. Following all injections into the MMN, labelled neurons were found in nuclei surrounding the mammillary body. The lateral and posterior subdivisions of the tuberomammillary nucleus projected mainly to the pars medianus and pars medialis of the MMN. The dorsal and ventral premammillary nuclei projected to the pars lateralis of the MMN. The supramammillary nucleus at rostral level had a small projection to the pars medialis and lateralis of the MMN. However, the most obvious projection from this nucleus was to the pars posterior of the MMN, chiefly from the lateral part of the caudal supramammillary nucleus. Injections into the lateral mammillary nucleus revealed inputs from the presubiculum, parasubiculum, septal region, dorsal tegmental nucleus, dorsal raphe nucleus, and periaqueductal gray. In addition, the lateral mammillary nucleus was found to receive a moderate projection from the medial part of the supramammillary nucleus and stronger projections from the lateral part of the caudal supramammillary nucleus. A very light projection was also seen from the lateral and posterior subdivisions of the tuberomammillary nucleus. These findings add to our knowledge of the extensive and complex connectivity of the mammillary nuclei. In particular, the local connections we have demonstrated with the supramammillary and tuberomammillary nuclei indicate the existence of significant local circuits as well as circuits involving more distant brain regions such as the septal nuclei, subiculum, prefrontal cortex, and brain stem tegmentum.

Afferent Pathways↗

Topographic organization of cortical and subcortical projections to posterior cingulate cortex in the cat: evidence for somatic, ocular, and complex subregions.

The posterior cingulate area (CGp) of the cat consists of cortex on the exposed cingulate gyrus and in the adjacent ventral bank of the splenial sulcus. We have placed deposits of distinguishable fluorescent tracers at multiple restricted sites in CGp and have analyzed the distribution throughout the forebrain of neurons labeled by retrograde transport. Cortical projections to CGp arise (in approximately descending order of strength) from anterior cingulate cortex; prefrontal cortex and premotor areas including the frontal eye fields; visual areas including especially areas 7 and 20b; parahippocampal areas; insular cortex; somesthetic areas; and auditory areas. Corticocortical pathways are organized topographically with respect to the posterior-anterior axis in CGp. Projections from prefrontal cortex and other areas with complex (as opposed to sensory, motor, or limbic) functions are concentrated posteriorly; projections from visual and oculomotor areas are concentrated at an intermediate level; and projections from areas with somesthetic and somatomotor functions are concentrated anteriorly. Thalamic projections to CGp arise from the anterior nuclei (AD, AV, and AM), from restricted portions of the ventral complex (VAd, VAm, and VMP), from discrete sectors of the lateral complex (LD, LPs, and LPm), from the rostral crescent of intralaminar nuclei (CM, PC, and CL), and from the reuniens nucleus. Projections from AM, VAd, LD, and LPs are spatially ordered in the sense that more ventral thalamic neurons project to more anterior cortical sites. Projections from AV and AD are stronger at more posterior cortical sites but do not show other signs of topographic ordering. Projections from LPm, CM, PC, CL, and RE are diffuse. We conclude (1) that cortical afferents of CGp derive predominantly from neocortical areas including those with well established sensory and motor functions; (2) that limbic projections to CGp originate primarily in structures, including the hippocampus, which are associated with memory, as opposed to structures, including the amygdala, which are associated with emotional and instinctual behavior; and (3) that CGp contains subregions in which complex, ocular, or somatic afferents predominate.

Afferent Pathways↗

Spinal projection neurons to the laterodorsal pontine tegmental nucleus: relationship to preganglionic neurons and nitric oxide synthase.

The region of the rat sacral parasympathetic nucleus (SPN) contains distinct subpopulations of neurons that project supraspinally or are preganglionic neurons. Some preganglionic neurons in the SPN serve as the motor outflow for urinary bladder contraction; other neurons in the SPN project to regions of the rostral pons that subserve micturition reflexes. Previous studies utilizing immunohistochemistry or staining for nicotinamide adenine dinucleotide phosphate-diaphorase (NADPH-d) have demonstrated that numerous neurons in the SPN contain nitric oxide synthase (NOS), the enzyme for nitric oxide synthesis. Thus, the objectives of this study were to determine 1) the distribution of neurons in the region of the SPN that project to the laterodorsal tegmentum (LDT) of the pons, 2) whether spinal neurons projecting to a peripheral autonomic ganglion also project to the LDT, and 3) whether NOS or NADPH-d is present in LDT projection neurons. Preganglionic neurons were identified by injecting the retrograde tracer fluorogold (FG) into the major pelvic ganglion (MPG). Supraspinally projecting neurons were identified by injecting the retrograde tracer fast blue (FB) into the LDT. Numerous FB-labeled neurons were present in the ipsi- and contralateral SPN and were immediately dorsal to FG-labeled preganglionic neurons. Neurons containing both tracers were not observed. Approximately 20% of preganglionic neurons, but no LDT projection neurons, were reactive for NOS and NADPH-d. These data suggest that the region of the SPN is a site for distinct subpopulations of neurons that project to the LDT and to the MPG and that NOS is contained in some preganglionic neurons, but is not a marker for LDT projection neurons.

Amidines↗

Development of ectopic projections from the ventral cochlear nucleus to the superior olivary complex induced by neonatal ablation of the contralateral cochlea.

The ability of an animal to localize a sound in space requires the precise innervation of the superior olivary complex by the ventral cochlear nuclei on each side of the lower brainstem. This precise pattern of innervation could require an immutable recognition of appropriate targets by afferent processes arising from these nuclei. This possibility was investigated by destroying one cochlea of gerbil pups (Meriones unguiculatus) on the second postnatal day and assessing the projections from the ventral cochlear nucleus (VCN) on the unablated side to the superior olivary complex during the subsequent 2 weeks and after the animals had reached maturity. A crystal of 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate (DiI) was inserted into VCN on the unablated side in animals ranging in age from 3 to 14 days. To assess the permanence of any altered pattern of innervation, horseradish peroxidase was injected into VCN on the unablated side in adult, neonatally ablated animals. Finally, electrophysiological responses to acoustic stimuli delivered to the ear on the unablated side were recorded in the superior olivary complex of adult animals to assess whether altered innervation patterns were functional. Normative data were derived from our accompanying study of the development of VCN projections to the superior olivary complex in normal gerbils (Kil et al., this issue). Whereas VCN normally projects to the lateral aspect of the ipsilateral medial superior olive and to the medial aspect of the contralateral medial superior olive in control animals, in experimental animals VCN on the unablated side projects to both sides of these nuclei. Whereas in the gerbil, VCN normally projects only to the hilar area and to the ventrolateral limb of the contralateral lateral superior olive, in experimental animals VCN on the unablated side projects throughout this nucleus. This induced projection is specific in that the efferents to each limb of the contralateral nucleus are linked to the normal projection to the homotopic region of the ipsilateral nucleus. Whereas VCN innervates the contralateral medial nucleus of the trapezoid body in control animals, in experimental animals VCN on the unablated side provides calyces of Held in the ipsilateral nucleus as well. The induced projections to these three major subnuclei of the superior olivary complex first appear within 24 hours of the cochlear ablation and continue to develop over at least the subsequent 11 days. Thus, prior to the day when the cochlea becomes functional, VCN has established specific ectopic projections to loci normally innervated by VCN on the ablated side.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Projections from the lateral, basal, and accessory basal nuclei of the amygdala to the entorhinal cortex in the macaque monkey.

We used the anterograde tracers Phaseolus vulgaris-leucoagglutinin (PHA-L) and biotinylated dextran amine (BDA) to examine the projections from the lateral, basal, and accessory basal nuclei of the amygdaloid complex to the entorhinal cortex in Macaca fascicularis monkeys. The heaviest amygdaloid projections originate in the lateral nucleus, which innervates the rostrally situated entorhinal fields but does not project to the caudal entorhinal cortex. The most extensive projections originate in the ventral division of the lateral nucleus. Injections in this subdivision lead to moderate to heavy fiber and terminal labeling in the entorhinal cortex, rostral levels of the rostral intermediate El (ER) and lateral fields, (ELr), and light labeling in the olfactory field EO. The projections from all portions of the lateral nucleus terminate most heavily in layer III. Layer II of EO and ER also receives a substantial input from the ventral division of the lateral nucleus. Layer II of ELr receives light innervation from all portions of the lateral nucleus that project to layer III. Projections from the basal nucleus arise mainly from the parvicellular division and are light to moderate in density. Fibers terminate predominantly in ELr, ER, EO, and the caudal portion of the lateral field (Elc); only the most rostral portion of El receives projections. While fibers from the basal nucleus innervate the same layers as the projections from the lateral nucleus, they tend to have a more vertical or radial orientation within the entorhinal cortex. Electron microscopic analysis of these fibers and terminals indicates that they overwhelmingly form asymmetrical synapses onto dendrites and dendritic spines. The accessory basal nucleus provides a light projection to the same regions of the entorhinal cortex innervated by the lateral and basal nuclei.

Amygdala↗

Organization of CA1 projections to the subiculum: a PHA-L analysis in the rat.

The organization of CA1 projections to the rat subiculum was investigated with the anterograde tracer, Phaseolus vulgaris leucoagglutinin (PHA-L). Discrete iontophoretic injections of PHA-L were placed into various transverse positions of the CA1 field at different septotemporal levels of the hippocampus. The distribution of CA1 projections was observed in dissected and extended hippocampal preparations. CA1 cells located proximally in the field, i.e., close to the CA2 field, gave rise to projections that terminated in the distal third of the subiculum, i.e., close to the presubiculum. CA1 cells located distally in the field, i.e., close to the subiculum, gave rise to projections that terminated proximally in the subiculum, i.e., just across the CA1/subiculum border. CA1 cells in the middle of the field projected to a midtransverse portion of the subiculum. The same general pattern of projections was observed at all septotemporal levels of the hippocampus. Varicose fibers from the CA1 neurons terminated among the basal dendrites of the subicular pyramidal cells, within the pyramidal cell layer, and in the deep portion of the molecular layer. In addition to the CA1 to subiculum projections, the discrete PHA-L injections provided the opportunity of examining the extent of local and associational connections within CA1. In general, associational connections in CA1 are far less extensive than in CA3. CA1 is not entirely without local connections, however. CA1 cells located close to the subicular border, for example, originated axons that first innervated the proximal subiculum and then reentered the CA1 field at the interface between stratum radiatum and stratum lacunosum-moleculare. In most of the experimental cases, there were collaterals located in stratum oriens of CA1 that branched from the fibers directed toward the subiculum. Thus, the basal dendrites of CA1 cells may receive associational inputs. The organization of the CA1 projections to the subiculum is discussed in relation to the organization of CA3 projections to CA1 and the differential output of transverse regions of the subiculum. The possibility is raised that information may be "channeled" through the hippocampal formation via the transverse organization of these connections and ultimately distributed to different recipients of hippocampal efferent projections.

Afferent Pathways↗

Differential estrogen accumulation among populations of projection neurons in the higher vocal center of male canaries.

The higher vocal center (HVC) of adult male canaries undergoes a seasonal change in volume that corresponds to seasonal modifications of vocal behavior: HVC is large when birds produce stereotyped song (spring) and is small when birds produce plastic song and add new song syllables into their vocal repertoires (fall). We reported previously that systemic exposure to testosterone (T) produces an increase in the volume of HVC similar to that observed with long-day photoperiods. T-induced growth of HVC occurred regardless of whether the borders of HVC were defined by Nissl-staining, the distribution of androgen-concentrating cells, or the distribution of projection neurons [separate neuronal populations within HVC project to the robust nucleus of the archistriatum (RA) and to Area X of the avian striatum (X)]. In the present study we used steroid autoradiography to determine whether T can influence the distribution of HVC cells that bind estrogen, and we combined estrogen autoradiography with retrograde labeling to determine whether HVC neurons that project to RA versus X differ in their ability to accumulate estrogen. Results showed that T increased the volume of Nissl-defined HVC and although HVC contained a low density of estrogen-concentrating cells, T increased the spatial distribution of these cells to match the Nissl borders of HVC. We also identified a region containing a high density of estrogen-concentrating cells located medial to HVC [we call this region paraHVC (pHVC)], and T also increased the volume of pHVC. pHVC also contained numerous X-projecting neurons, but few if any RA-projecting neurons. Double-labeling analysis revealed that RA-projecting neurons did not accumulate estrogen, a small percentage of X-projecting neurons in HVC accumulated estrogen, and the majority of X-projecting neurons in pHVC showed heavy accumulation of estrogen. The data reported here and in our previous article suggest distinct roles for gonadal steroids within the HVC-pHVC complex: estrogens are concentrated by neurons that project to a striatal region that influences vocal production during song learning (X), whereas androgens are concentrated primarily by neurons that project to a motor region that is involved in vocal production during both song learning and the recitation of already-learned song (RA).

Animals↗

Changes in the segmental pattern of sensory neuron projections in the chick hindlimb under conditions of altered cell adhesion molecule function.

In the developing chick hindlimb, pathfinding by sensory axons is affected by their interactions with other axons entering the limb. Cell adhesion molecules (CAMs) expressed on the growing axons are likely to influence these interactions. Accordingly, to elucidate how CAMs affect sensory axon pathfinding, we injected antibodies that block the functions of NCAM, G4/L1, or N-cadherin into the hindlimb, starting at St. 25, when all motoneuron axons but only a few sensory axons had entered the plexus. In each case, the segmental pattern of projections was assessed 2-3 days later by retrogradely labeling individual peripheral nerves. With all perturbations, the gross anatomical nerve pattern developed normally; that is, some sensory axons formed cutaneous nerves while other sensory axons projected to muscles. However, the segmental pattern of sensory projections was changed when either G4/L1 or N-cadherin function was blocked in that fewer sensory axons crossed the anterior-posterior axis of the plexus. A likely reason for this effect is that anti-G4/L1 and anti-N-cadherin each decreased the amount of fasciculation and that sensory axons are less able to travel across the plexus when they are defasciculated. Anti-G4/L1 affected both cutaneous and muscle sensory projections while anti-N-cadherin affected cutaneous but not muscle sensory projections, in accord with known differences in the expression of these two CAMs on sensory and motoneuron axons. Although anti-NCAM did not appear to alter sensory projections, when polysialic acid (PSA) was enzymatically removed from NCAM, there was a marked increase in cutaneous projections from the most proximate DRG, although muscle sensory projections were unchanged. PSA removal may cause an increase in fasciculation that forces sensory axons to track along neighboring axons. Thus, without PSA, cutaneous axons project more in accord with the relative anterior-posterior positions they had as they entered the plexus. Taken together, these studies suggest that axonal fasciculation mediated by CAMs and regulated by PSA influences the ability of sensory growth cones to navigate through the plexus and project along the correct peripheral nerves.

Afferent Pathways↗