Identification of the reptilian basolateral amygdala: an anatomical investigation of the afferents to the posterior dorsal ventricular ridge of the lizard Podarcis hispanica

The presence of multimodal association in the telencephalon of reptiles has been investigated by tracing the afferent connections to the posterior dorsal ventricular ridge (PDVR) of the lizard Podarcis hispanica. The PDVR receives telencephalic afferents from the lateral (olfactory) and dorsal cortices, and from the three unimodal areas of the anterior dorsal ventricular ridge, in a convergent manner. From the diencephalon, it receives afferents from the dorsomedial anterior and medial posterior thalamic nuclei, and from several hypothalamic nuclei. Brainstem afferents to the PDVR originate in the dorsal interpeduncular nucleus, the nucleus of the lateral lemniscus and parabrachial nucleus. The afferents to the thalamic nuclei that project to the PDVR have also been studied. The dorsomedial anterior thalamic nucleus receives projections mainly from limbic structures, whereas the medial posterior thalamic nucleus is the target of projections from structures with a clear sensory significance (optic tectum, torus semicircularis, nuclei of the lateral and spinal lemniscus, superior olive and trigeminal complex). As a result, the PDVR appears as an associative centre that receives visual, auditory, somatosensory and olfactory information from several telencephalic and non-telencephalic centres, and a multimodal projection from the medial posterior thalamic nucleus. This pattern of afferents of the PDVR is similar to that of the caudal neostriatum in birds and the basolateral division of the mammalian amygdala. These results indicate that a multimodal amygdala is already present in reptiles, and has probably played a key role in the evolution of the vertebrate brain.     

Afferent and efferent connections of the bullfrog medial pallium.

Horseradish peroxidase or tritiated proline was unilaterally injected into the medial pallium in bullfrogs in order to determine the sources of afferent projections to the medial pallium and the targets of pallial efferent projections. Some cells in all telencephalic centers, except the corpus striatum and the pars lateralis of the amygdala, project to the ipsilateral medial pallium. The medial pallium receives projections from fewer centers in the contralateral hemisphere, which include the medial septal nucleus, the pars medialis of the amygdala, the bed nucleus of the pallial commissure and the medial pallium. The raphe nucleus and the anterior thalamic nuclei appear to be the only sources of afferents to the medial pallium from outside the telencephalon. Efferents of the medial pallium are far more extensive than reported in earlier studies. The medial pallium projects ipsilaterally to all telencephalic nuclei, with the exception of a large part of the corpus striatum, and contralaterally to the medial septal nucleus, the olfactory tubercle, amygdala, medial pallium and bed nucleus of the pallial commissure. Extensive efferent projections also terminate in preoptic and hypothalamic regions, as well as in most thalamic relay nuclei, the pretectum and, possibly, the optic tectum. Similarities to the medial pallium in other tetrapods and to that in mammals suggest that the medial pallium in anurans is homologous to the subicular and CA fields and, possibly, the dentate gyrus in mammals. However, the extensive projections of the medial pallium to the dorsal thalamus and pretectum in anurans may be primitive features of the medial pallium retained in anurans, or uniquely derived features in anurans.     

Connections of the ventral telencephalon (subpallium) in the zebrafish (Danio rerio)

Connections of the medial precommissural subpallial ventral telencephalon, i.e., dorsal (Vd, interpreted as part of striatum) and ventral (Vv, interpreted as part of septum) nuclei of area ventralis telencephali, were studied in the zebrafish (Danio rerio) using two tracer substances (DiI or biocytin). The following major afferent nuclei to Vd/Vv were identified: medial and posterior pallial zones of dorsal telencephalic area, and the subpallial supracommissural and postcommissural nuclei of the ventral telencephalic area, the olfactory bulb, dorsal entopeduncular, anterior and posterior parvocellular preoptic and suprachiasmatic nuclei, anterior, dorsal and central posterior dorsal thalamic, as well as rostrolateral nuclei, periventricular nucleus of the posterior tuberculum, posterior tuberal nucleus, various tuberal hypothalamic nuclei, dorsal tegmental nucleus, superior reticular nucleus, locus coeruleus, and superior raphe nucleus. Efferent projections of the ventral telencephalon terminate in the supracommissural nucleus of area ventralis telencephali, the posterior zone of area dorsalis telencephali, habenula, periventricular pretectum, paracommissural nucleus, posterior dorsal thalamus, preoptic region, midline posterior tuberculum (especially the area dorsal to the posterior tuberal nucleus), tuberal (midline) hypothalamus and interpeduncular nucleus. Strong reciprocal interconnections likely exist between septum and preoptic region/midline hypothalamus and between striatum and dorsal thalamus (dopaminergic) posterior tuberculum. Regarding ascending activating/modulatory systems, the pallium shares with the subpallium inputs from the (noradrenergic) locus coeruleus, and the (serotoninergic) superior raphe, while the subpallium additionally receives such inputs from the (dopaminergic) posterior tuberculum, the (putative cholinergic) superior reticular nucleus, and the (putative histaminergic) caudal hypothamalic zone.     

Noradrenergic projections to the song control nucleus area X of the medial striatum in male zebra finches (Taeniopygia guttata)

There is considerable functional evidence implicating norepinephrine in modulating activity in the vocal control circuit of songbirds. However, our knowledge of noradrenergic inputs to the song system is incomplete. In this study, cholera toxin subunit B (CTB) injections into area X revealed projections from the noradrenergic nuclei locus coeruleus and subcoeruleus, and injections of biotinylated dextran amines into these noradrenergic nuclei labeled fibers in area X. The nonreciprocity of this connection was demonstrated by the absence of retrogradely labeled cells in area X following injections of CTB into the locus coeruleus. Additionally, we found novel inputs to area X from the nidopallium and arcopallium, the mesencephalic central gray, and the dorsolateralis anterior (DLL) and posterior (DLP) lateralis in the thalamus. Area X can be clearly distinguished from the surrounding medial striatum based on cytoarchitectural and chemical neuroanatomical criteria. We show here that neuromodulatory inputs to area X however, exhibit a considerable degree of overlap with the surrounding area. This finding suggests that regional specificity in neuromodulator action is most likely afforded by a specialization in receptor density and enzyme distribution rather than projections from the synthesizing nuclei. Our results extend current knowledge about noradrenergic projections to specialized nuclei of the song control circuit and provide neuroanatomical evidence for the functional action of norepinephrine-modulating context-dependent ZENK expression in area X. Furthermore, the novel projections to area X from telencephalic and thalamic areas could be new and interesting nodes in the striatopallidothalamic loop spanning the songbird brain.     

Unilateral ablation of telencephalon induces appearance of contralateral cortical and subcortical projections to thalamic nuclei

This study sought to investigate the afferent connections of the thalamus in the rat following massive telencephalic lesions. After unilateral removal of the telencephalon, horseradish peroxidase (HRP) was injected into the thalamic nuclei ipsilateral or, for control purposes, contralateral to the lesioned side. Injection of HRP into the ventral or posterior thalamus ipsilateral to the lesioned telencephalon led to retrograde transport and HRP-labeling of cells in unilateral projection areas in the cortex, diencephalon and mesencephalon and, in addition, unexpectedly, in their mirror-image sites in the hemisphere contralateral to the injected thalamus. HRP-positive cells in the contralateral hemisphere were found whenever HRP was injected 7 days postlesion, when the animals had ceased to exhibit spontaneous turning behavior, but not when it was injected immediately after the lesion. We favor the hypothesis that the development of contralateral afferents to the thalamic nuclei represents a morphological substrate of the behavioral reorganization necessitated by the asymmetrical removal of the telencephalon.     

Connections of some auditory-responsive posterior thalamic nuclei putatively involved in activation of the hypothalamo-pituitary-adrenocortical axis in response to audiogenic stress in rats: an anterograde and retrograde tract tracing study combined with Fos expression

Prior studies in our laboratory demonstrated that part of the thalamus is necessary for activating the hypothalamo-pituitary-adrenocortical (HPA) axis in response to audiogenic stress in rats. The present studies were designed to determine how the auditory-responsive thalamic nuclei might activate the HPA axis. Both retrograde [Fluoro-Gold (FG)] and anterograde [Phasoleus vulgaris-leucoagglutinin (PHA-L) and biotinylated dextran amines (BDA)] tracers were employed to study the putative connectivity between the thalamus and the medial parvocellular region of the hypothalamic paraventricular nucleus (PAmp). In addition, rats receiving FG in the PAmp were subjected to audiogenic stress, and the distribution of both FG and the protein product of the immediate-early gene c-fos, Fos, were determined by double immunohistochemistry, to help assess putative functional links between the auditory-responsive thalamic nuclei and PAmp. The results of PAmp FG placement indicated retrogradely labeled cells in several areas, including the bed nucleus of the stria terminalis, hypothalamic regions, the supramammillary nucleus, some thalamic regions, and importantly, a few multisensory nuclei of the thalamus, including the parvicellular division of the subparafascicular and posterior intralaminar nuclei. Injections of the tracers PHA-L or BDA into these auditory-responsive posterior thalamic nuclei provided further evidence of projections to the PAmp. In addition, several forebrain areas were observed to receive moderate to heavy innervation. These areas included most of the regions described above, which, in turn, project to the PAmp. Because cells in the multisensory thalamic nuclei, hypothalamic, and forebrain areas were double labeled with FG and Fos, the results suggest that either direct projections from the thalamus to PAmp neurons, or indirect projections from the thalamus to stress-responsive forebrain areas projecting to the PAmp, might mediate activation of the HPA axis by audiogenic stress.     

Efferent projections of the ectostriatum in the pigeon (Columba livia)

The ectostriatum is a major visual component of the avian telencephalon. The core region of the ectostriatum (Ec) receives visual input from the optic tectum through thalamic nuclei. In the present study, the efferent projections of the ectostriatum were investigated by using the anterograde tracers Phaseolus vulgaris leucoagglutinin and biotinylated dextran amine. Projection patterns resulting from these tracers were confirmed by the retrograde tracer cholera toxin subunit B. When anterograde tracers were injected in Ec, primary projections were seen traveling dorsolaterally to the belt region of the ectostriatum (Ep) and the neostriatal area immediately surrounding Ep (Ep2). Neurons in Ep sent projections primarily to the overlying Ep2. The efferents of Ep2 traveled dorsolaterally to terminate in three telencephalic regions, from anterior to posterior: (1) neostriatum frontale, pars lateralis (NFL), (2) area temporo-parieto-occipitalis (TPO), and (3) neostriatum intermedium, pars lateralis (NIL). A part of the archistriatum intermedium and the lateral part of the neostriatum caudale also received somewhat minor projections. In addition, some neurons in Ec were also the source of direct, but minor, projections to the NFL, TPO, NIL, and archistriatum intermedium. The topographical relationship among the primary (Ec), secondary (Ep and Ep2), and tertiary (NFL, TPO, NIL) areas indicate that the neural populations for visual processing are organized along the rostral-caudal axis. Thus, the anterior Ec sent efferents to the anterior Ep, which in turn sent projections to anterior Ep2. Neurons in the anterior Ep2 sent projections to NFL and the anterior TPO. Similarly, the intermediate and posterior Ec sent projections to corresponding parts of Ep, whose efferents projected to intermediate and posterior Ep2, respectively. The intermediate Ep2 gave rise to major projections to TPO, whereas posterior Ep2 neurons sent efferents primarily to NIL. The organization of this neural circuit is compared with those of other sensory circuits in the avian telencephalon, as well as the laminar arrangement of the mammalian isocortex.     

Organization of thalamic afferents to anterior dorsal ventricular ridge in turtles. I. Projections of thalamic nuclei

Dorsal ventricular ridge (DVR) is a thalamorecipient, subcortical telencephalic structure in reptiles and birds. Although there is a fair amount of information about sources of afferents to DVR, little is known about the relationship of projections from individual thalamic nuclei to the organization of the structure. This study examines the relationship between thalamic projections and both areal and zonal divisions of anterior DVR (ADVR; Balaban,'78a) of emydid turtles with orthograde degeneration, and horseradish peroxidase techniques. Individual thalamic nuclei contribute either a diffuse or a restricted projection to ADVR. Diffuse projections arise primarily from the dorsomedial anterior nucleus. These fine-caliber axons distribute bilaterally over a wide region of the telencephalon via both medial and lateral thalamotelencephalic pathways. The terminal regions include septum, striatum and the medial bank of cortex caudal to the lamina terminalis. In ADVR, the fibers are distributed sparsely in zones 2–4 of dorsal, medial and ventral areas. Restricted projections to ADVR originate in nucleus rotundus, nucleus reuniens and nucleus caudalis. They ascend ipsilaterally in the lateral thalamotelencephalic pathway (lateral forebrain bundle), and enter ADVR rostral to the anterior commissure. Nucleus rotundus projects to zone 4 of dorsal area, nucleus caudalis projects to zones 2–4 of the dorsal division of medial area, and nucleus reuniens projects to zones 2–4 of both the ventral division of medial area and the ventral area. Comparison of these results with thalamotelencephalic projections in mammals suggests that diffuse and restricted thalamic projection systems are a common feature of both groups. Restricted thalamic projections in reptiles, birds and mammals, terminating in anatomically distinct regions, also appear to be associated with different sensory modalities. The significance of diffuse systems is not clear.     

Forebrain connections of the gustatory system in ictalurid catfishes

Horseradish peroxidase tracing and extracellular electrophysiological recording techniques were employed to delineate prosencephalic connections of the gustatory system in ictalurid catfishes. The isthmic secondary gustatory nucleus projects rostrally to several areas of the ventral diencephalon including the nucleus lobobulbaris and the nucleus lateralis thalami. Injections of HRP in the vicinity of the nucleus lobobulbaris reveal an ascending projection to the telencephalon terminating in the area dorsalis pars medialis (Dm) and the medial region of area dorsalis pars centralis (Dc). Conversely, injections of HRP into the gustatory region of area dorsalis pars medialis label small neurons in the nucleus lobobulbaris. Gustatory neurons in the telencephalon send descending projections via the medial and lateral forebrain bundles to several nuclei in the anterior and ventroposterior diencephalon. The nucleus lateralis thalami, a diencephalic nucleus, receives ascending gustatory projections from the secondary gustatory nucleus but does not project to the telencephalon. Neurons in both the nucleus lateralis thalami and the telencephalic gustatory target exhibit multiple extraoral and oral receptive fields and complex responses to chemical (taste) and tactile stimulation.     

鸣禽中脑听觉核团边缘区神经投射的研究

对鸣禽白腰文鸟（Ｌｏｎｃｈｕｒａ ｓｔｒｉａｔａ）中脑听觉核团－背外侧核（ｎｕｃｌｅｕｓ ｍｅｓｅｎｃｅｐｈａｌｉｃｕｓ ｌａｔｅｒａｌｉｓ,ｐａｒｓ ｄｏｒｓａｌｉｓ,ＭＬｄ）和丘间核（ｎｕｃｌｅｕｓ ｉｎｔｅｒｃｏｌｌｉｃｕｌａｒｉｓ,ＩＣｏ）的脑啡肽免疫化学特异和神经联系进行了研究,结果发现：脑啡肽能标记纤维或细胞主要分布于中脑听觉核团的痛外侧核边缘区、丘间核,而背外侧核中央几乎无分布。双向     

The primate mediodorsal (MD) nucleus and its projection to the frontal lobe

The frontal lobe projections of the mediodorsal (MD) nucleus of the thalamus were examined in rhesus monkey by transport of retrograde markers injected into one of nine cytoarchitectonic regions (Walker's areas 6, 8A, 9, 10, 11, 12, 13, 46, and Brodmann's area 4) located in the rostral third of the cerebrum. Each area of prefrontal, premotor, or motor cortex injected was found to receive a topographically unique thalamic input from clusters of cells in specific subdivisions within MD. All of the prefrontal areas examined also receive topographically organized inputs from other thalamic nuclei including, most prominently, the ventral anterior (VA) and medial pulvinar nuclei. Conversely, and in agreement with previous findings, MD projects to areas of the frontal lobe beyond the traditional borders of prefrontal cortex, such as the anterior cingulate and supplementary motor cortex. The topography of thalamocortical neurons revealed in coronal sections through VA, MD, and pulvinar is circumferential. In the medial part of MD, for example, thalamocortical neurons shift from a dorsal to a ventral position for cortical targets lying medial to lateral along the ventral surface of the lobe; neurons in the lateral MD move from a ventral to a dorsal position, for cortical areas situated lateral to medial on the convexity of the hemisphere. The aggregate evidence for topographic specificity is supported further by experiments in which different fluorescent dyes were placed in multiple areas of the frontal lobe in each of three cases. The results show that very few, if any, thalamic neurons project to more than one area of cortex. The widespread cortical targets of MD neurons together with evidence for multiple thalamic inputs to prefrontal areas support a revision of the classical hodological definition of prefrontal cortex as the exclusive cortical recipient of MD projections. Rather, the prefrontal cortex is defined by multiple specific relationships with the thalamus.     

Comparison of cerebellothalamic and pallidothalamic projections in the monkey (Macaca fuscata): A double anterograde labeling study

To address the question of segregated projections from the internal segment of the globus pallidus (GPi) and the cerebellar nuclei (Cb) to the thalamus in the monkey, we employed a double anterograde labeling strategy combining the anterograde transport of horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP) with biotinylated dextran amine (BDA) transport. The tissue was processed sequentially for WGA-HRP, and then BDA immunohistochemistry using two different chromogens. Since the two labels were easily distinguishable on the same histological section, the interrelationship between the cerebellar and pallidal projection systems could be directly evaluated. We found that both the cerebellothalamic and pallidothalamic label consisted of dense plexuses of labeled fibers and swellings in a patch-like configuration. The patches or foci of labeling were distributed either as dense single label or as interdigitating patches of double label. We found dense single label in the central portion of the ventral anterior nucleus pars principalis (VApc) and the ventral lateral nucleus pars oralis (VLo) following the GPi injections or in the central portion of the ventral posterior lateral nucleus pars oralis (VPLo) and nucleus X (X) following the cerebellar nuclei injections. Complementary interdigitating patches of WGA-HRP and BDA labeling were found primarily in transitional border regions between thalamic nuclei. On occasion, we found overlap of both labels. We observed a gradient pattern in the density of the pallidothalamic and cerebellothalamic projections. The pallidothalamic territory included VApc, VLo, and the ventral lateral nucleus pars caudalis (VLc), with the density of these projections decreasing along an anterior to posterior gradient in the thalamus. Occasional patches of pallidal label were found in VPLo and nucleus X. Conversely, the density of cerebellothalamic projections increased along the same gradient, with the cerebellothalamic territory extending anteriorly beyond the cell-sparse zones of VPLo, X, and VLc to include VLo and VApc also. These data suggest that although the cerebellar and pallidal projections primarily occupy separate thalamic territories, individual thalamic nuclei receive differentially weighted inputs from these sources. © 1996 Wiley-Liss, Inc.     

Nuclear organization of the bullfrog diencephalon

A cytoarchitectonic analysis was performed on the diencephalic nuclei of the bullfrog, Rana catesbeiana. The epithalamus contains two widely recognized habenular nuclei. The thalamus has three subdivisions: dorsal and ventral thalamus, and posterior tuberculum. The dorsal thalamus may be further parcelled into anterior, middle, and posterior zones. Connectional data from other studies support this zonation. The anterior zone projects to the telencephalic pallium. The middle zone nuclei receive a strong input from the midbrain roof and project to the telencephalic striatal complex. The posterior zone nuclei do not appear to project to the telencephalon; they may eventually be placed in the pretectum, a transitional area between the diencephalon and mesencephalon. Two of the ventral thalamic populations have been frequently placed in the dorsal thalamus and called the nucleus rotundus and the lateral geniculate nucleus. These terms imply homology with sauropsid dorsal thalamic nuclei, but our analysis and current connectional information do not support such homologies. We have given these populations more neutral names. The hypothalamus is divisible into a preoptic and infundibular hypothalamus, and the preoptic area can be further separated into anterior and posterior preoptic areas. The posterior area contains the magnocellular preoptic nucleus and a dorsal arm of this nucleus, often placed in the ventral thalamus, was recognized. We have tentatively placed the posterior entopeduncular nucleus in the hypothalamus.     

The origin of projections from the posterior cingulate and retrosplenial cortices to the anterior,medial dorsal and laterodorsal thalamic nuclei of macaque monkeys

Interactions between the posterior cingulate cortex (areas 23 and 31) and the retrosplenial cortex (areas 29 and 30) with the anterior, laterodorsal and dorsal medial thalamic nuclei are thought to support various aspects of cognition, including memory and spatial processing. To detail these interactions better, the present study used retrograde tracers to reveal the origins of the corticothalamic projections in two closely related monkey species (Macaca mulatta, Macaca fascicularis). The medial dorsal thalamic nucleus received only light cortical inputs, which predominantly arose from area 23. Efferents to the anterior medial thalamic nucleus also arose principally from area 23, but these projections proved more numerous than those to the medial dorsal nucleus and also involved additional inputs from areas 29 and 30. The anterior ventral and laterodorsal thalamic nuclei had similar sources of inputs from the posterior cingulate and retrosplenial cortices. For both nuclei, the densest projections arose from areas 29 and 30, with numbers of thalamic inputs often decreasing when going dorsal from area 23a to 23c and to area 31. In all cases, the corticothalamic projections almost always arose from the deepest cortical layer. The different profiles of inputs to the anterior medial and anterior ventral thalamic nuclei reinforce other anatomical and electrophysiological findings suggesting that these adjacent thalamic nuclei serve different, but complementary, functions supporting memory. While the lack of retrosplenial connections singled out the medial dorsal nucleus, the very similar connection patterns shown by the anterior ventral and laterodorsal nuclei point to common roles in cognition.     

Biotinylated dextran amine as an anterograde tracer for single- and double-labeling studies.

Fluorescent dextran amines have recently been reported to be useful for anterograde pathway tracing. However, fluorescent markers are not always ideal for detailed mapping studies. We therefore evaluated the efficacy of a biotinylated dextran amine (BDA) for anterograde labeling in several different preparations. BDA was visualized with an avidin-biotinylated HRP (ABC) procedure followed by a standard or metal-enhanced diaminobenzidine (DAB) reaction. After iontophoretic injections of BDA into neocortex-like telencephalic regions in pigeons or into visual or somatosensory cortex in rats, there was excellent and abundant labeling of axons and terminals in forebrain, midbrain and hindbrain target areas with 1-week survival times. Large pressure injections of BDA into the avian telencephalon were also found to result in extensive anterograde labeling. We then carried out a series of studies using 2-color DAB double-labeling to determine effective approaches for combining BDA labeling with other labeling methods. Using an isolated embryonic chick spinal cord-hindlimb preparation, we combined BDA labeling with another anterograde labeling method to differentially label two sets of projections. In these studies, sensory neuron and motoneuron projections into the limb from the same segmental level, or motoneuron projections into the limb from two separate segments were differentially labeled by using HRP (visualized first with a blue/black metal-DAB reaction) and BDA (visualized second with a brown DAB reaction). In other double-labeling studies, we combined BDA labeling of axons and terminals with immunohistochemical labeling of neurons. In these experiments, telencephalic neurons in pigeons or rats were labeled immunohistochemically for parvalbumin or substance P (using a brown DAB reaction) and BDA-labeled axons were labeled blue/black (using a metal-intensified DAB reaction). Double-labeling was successful regardless of whether the entire immunohistochemical labeling procedure preceded or followed the BDA labeling procedure. Together, these studies show that BDA is effective for anterograde pathway tracing and can be used in double-label studies with other labeling methods.     

Telencephalic afferent nuclei in the Carp diencephalon, with special reference to fiber connections of the nucleus preglomerulosus pars lateralis

Fiber connections of the nucleus preglomerulosus pars lateralis (PGl), which primarily provides afferent inputs to the telencephalon, were examined in carp by means of horseradish peroxidase tracing methods. The major afferent sources of PGl are the bilateral nucleus tuberis anterior and a few projections are found deriving from the ipsilateral nucleus ventromedialis thalami, nucleus centralis posterior, dorsal periventricular hypothalamus, and torus semicircularis. Axons arising in the PGl can be traced to the area dorsalis pars centralis of the ipsilateral telencephalon, nucleus centralis posterior and the nucleus ventromedialis thalami. In addition, another 5 telencephalic afferent nuclei are found in the diencephalon; the nucleus subrotundus of Sheldon, nucleus preglomerulosus pars anterior, nucleus ventromedialis thalami, nucleus centralis posterior and nucleus posterior thalami.     

Telencephalic projections from midbrain and isthmal cell groups in the pigeon. I. Locus coeruleus and subcoeruleus

Horseradish peroxidase (HRP) and amino acid autoradiography were used in pigeon to determine the trajectories and projection patterns of neurons within the locus coeruleus and subcoeruleus nuclei upon the cerebral hemispheres. The specific cell groups investigated include the locus coeruleus (LoC), nucleus subcoeruleus dorsalis, and nucleus subcoeruleus ventralis. Efferents from each of these nuclei ascend to the telencephalon via the medial and lateral forebrain bundles, ansa lenticularis, and the quintofrontal and occipitomesencephalic tracts. A separate dorsally situated bundle derived from LoC neurons reaches many dorsal thalamic nuclei. The telencephalic projections of the LoC and subcoeruleus nuclei are bilateral and symmetrical, although projections to the contralateral hemisphere are sparse. Crossing fibers project to contralateral targets primarily via the dorsal supraoptic decussation and along the dorsal and ventral margins of the anterior commissure. Within the telencephalon, the following neural structures receive input from neurons in the LoC and subcoeruleus cell groups: the paleostriatal complex including the paleostriatum augmentatum and lobus parolfactorius, septal nuclei, nucleus accumbens, olfactory tubercle, hippocampus and parahippocampal area, nucleus taeniae, dorsal archistriatum, lateral neostriatum, hyperstriatum dorsale, hyperstriatum ventrale, and preoptic area. Large portions of the cerebral hemispheres including the hyperstriatum accessorium, much of the neostriatum and hyperstriatum ventrale, and all but the dorsal portion of the archistriatum receive little or no input from either the locus coeruleus or subcoeruleus cell groups. This is apparently different from the condition in mammals in which virtually all cortical fields receive input from neurons within the LoC. Moreover, the pattern of projections of the subcoeruleus nuclei upon telencephalic fields described here as well as recent histochemical data suggest that these cell groups are comparable to the lateral tegmental (A8) cell group of mammals rather than to the mammalian subcoeruleus nuclei.     

The avian somatosensory system: connections of regions of body representation in the forebrain of the pigeon

In order to establish the basic connectivity of physiologically identified somatosensory regions of the thalamus and telencephalon in the pigeon, injections of wheatgerm agglutinin-horseradish peroxidase were made under electrophysiological control and the projections were charted following conventional neurohistochemistry. The physiological recordings generally confirmed the findings of Delius and Bennetto (Brain Research, 37 (1972) 205-221) of somatosensory sites within the dorsal thalamus, anterior hyperstriatum and caudomedial neostriatum, and the anatomical results show that the thalamic cells of origin of the projections to the two telencephalic regions are largely separate: a rostral cell group comprising nucleus dorsalis intermedius ventralis anterior projects to the anterior hyperstriatum accessorium (HA), whilst a caudal cell group comprising caudal regions of nucleus dorsolateralis posterior (DLP) projects to the medial neostriatum intermedium and caudale (NI/NC). Caudal DLP is also the origin of a visual projection to NI/NC, and its terminal field also approximates that of the thalamic auditory nucleus ovoidalis. Since the anterior HA and NI/NC were here shown to be reciprocally connected, there is a possibility of multimodal input to both telencephalic regions. HA was also further defined as the origin of the basal branch of the septomesencephalic tract, and hence potentially provides an outlet for both telencephalic somatosensory regions. The results are discussed within a comparative context.     

Corticothalamic projections from layer 5 of the vibrissal barrel cortex in the rat

This study bears on the projections of layer 5 cells of the vibrissal sensory cortex to the somatosensory thalamus in rats. Small groups of cells were labeled with biotinylated dextran amine (BDA), and their axonal arborizations were individually reconstructed from horizontal sections counterstained for cytochrome oxidase. Results show that the vast majority ( approximately 95%) of layer 5 axons that innervate the somatosensory thalamus are collaterals of corticofugal fibers that project to the brainstem. The anterior pretectal nucleus, the deep layers of the superior colliculus, and the pontine nuclei are among the structures most often coinnervated. In the thalamus, layer 5 axons terminate exclusively in the dorsal part of the posterior group (Po), where they form clusters of large terminations. Because dorsal Po projects to multiple cortical areas, we sought to determine whether all recipient areas return a layer 5 projection to this part of the thalamus. Additional experiments using fluoro-gold and BDA injections provided evidence that the primary somatosensory area is the sole source of layer 5 projections to dorsal Po but that this thalamic region receives convergent layer 6 projections from the primary and second somatosensory areas and from the motor and insular cortices. These results show that layer 5 projections do not overlap in associative thalamic nuclei, thus defining area-related subdivisions. Furthermore, the coinnervation of brainstem nuclei by layer 5 CT axons suggests that this pathway conveys to the thalamus a copy of the cortical output aimed at brainstem structures.