Topographic organization of subcortical projections to the anterior thalamic nuclei in the rat.

Subcortical projections to the anterior thalamic nuclei were studied in the rat, with special reference to projections from the mammillary nuclei, by retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase. The medial mammillary nucleus (MM) projects predominantly ipsilaterally to the entire anterior thalamic nuclei, whereas the lateral mammillary nucleus projects bilaterally to the anterodorsal nucleus (AD) of the anterior thalamic nuclei. A topographic relationship was recognized between the MM and the anterior thalamic nuclei. The dorsal region of the pars mediana of the MM projects to the interanteromedial nucleus (IAM), whereas the ventral region projects to the rostral part of the anteromedial nucleus (AM). The dorsal and the ventral regions of the pars medialis project to the dorsomedial part of the AM at its caudal and rostral levels, respectively. The dorsomedial region of the pars lateralis projects to the ventral AM. The ventrolateral region of the pars lateralis projects to the ventral part of the anteroventral nucleus (AV) in such a manner that rostral cells project rostrally and caudal cells project caudally. The pars basalis projects predominantly ipsilaterally to the dorsolateral AV and bilaterally to the AD. The rostrolateral region of the pars posterior projects to the lateral AV, whereas the medial and the caudal regions of the pars posterior project to the dorsomedial AV. The rostrodorsal part of the nucleus reticularis thalami was found to project to the anterior thalamic nuclei; cells located rostrally in this part project to the IAM and AM, whereas cells located caudodorsally project to the AV and AD. The laterodorsal tegmental nucleus projects predominantly ipsilaterally to the AV, especially to its dorsolateral part. The present study demonstrates that subdivisions of the subcortical structures are connected to the subnuclei of the anterior thalamic nuclei, with a clear-cut topography arranged in the dorsoventral and the rostrocaudal dimensions.     

Organization of connections between the thalamic reticular and the anterior thalamic nuclei in the rat

The thalamic reticular nucleus (TRN) receives topographically organized input from specific sensory nuclei such as the lateral geniculate nucleus. The present study shows this in the rat. However, the pattern of thalamic connections to the limbic reticular sector is unknown. Injecting biocytin into the ventral parts of anteroventral and anteromedial nuclei labeled neurons and axons in the rostral TRN. Filled axon collaterals and their terminals occupied a rectangular sheet in a plane close to the horizontal, and were confined to the inner zone (the medial portion) of the limbic TRN. Retrogradely filled cells were in the middle of the rostral pole in the same horizontal plane, receiving synapses from surrounding labeled boutons. In electron micrographs, thalamic terminals were found to contain round, densely packed synaptic vesicles and formed asymmetrical synapses onto reticular somata and dendritic profiles. Displacing the injection site along the dorso-ventral and rostro-caudal axs in the anterior nuclei produced corresponding shifts of antero- and retrograde labeling within the inner reticular zone. Projections from the dorsal portions of the anterior nuclei did not follow this pattern. Axons from the anterodorsal nucleus occupied the rostralmost tip of both inner and outer zones of the dorsal limbic sector. In accordance with earlier reports, the limbie sector was found to represent several dorsal thalamic nuclei parallel to each other medio-laterally. A topography is described for the limbic reticulo-thalamic connections, suggesting that the rostral TRN is able to influence circumscribed areas of the limbic thalamus. © 1995 Wiley-Liss, Inc.     

Cortical,thalamic, and amygdaloid projections of rat temporal cortex

The cortical, thalamic, and amygdaloid connections of the rodent temporal cortices were investigated by using the anterograde transport of iontophoretically injected biocytin. Injections into area Te1 labeled axons and terminals in the ventral regions of the dorsal and ventral subnuclei of the medial geniculate complex, area Te3, the rostrodorsal part of area Te2, and the ventrocaudal caudate putamen. No amygdaloid labeling was observed. Thalamic projections from Te2 targeted the lateral posterior nucleus, the dorsal part of the dorsal subnucleus of the medial geniculate complex, and the peripeduncular nucleus. Corticocortical projections mainly terminated in the dorsal perirhinal cortex, but moderately dense projections were observed in medial and lateral peristriate cortex, and only light projections were observed to Te1 and Te3. Projections to these isocortical regions terminated in layers I and VI. Amygdaloid projections targeted the ventromedial subdivision of the lateral nucleus and the adjacent part of the anterior basolateral nucleus. Area Te3 was observed to project to the ventrolateral parts of the dorsal and ventral subnuclei of the medial geniculate complex, the dorsal perirhinal cortex, rostral Te2, and Te1. In the amygdala, labeled fibers and terminals were concentrated in the dorsolateral subdivision of the lateral nucleus. These data confirm that areas Te1 and Te3 are hierarchically organized cortical areas connected with auditory relay nuclei in the thalamus. Area Te2, in contrast, appears to be weakly connected with Te1 and Te3 but is heavily connected with the peristriate cortex and tectorecipient thalamic nuclei. Te2 appears to be a visually related cortical area. The data also indicate that projections from Te2 and Te3 target different subregions of the lateral nucleus and that Te2, but not Te3, projects to the basolateral nucleus. J. Comp. Neurol. 382:153-175, 1997. © 1997 Wiley-Liss, Inc.     

Anterior thalamic afferents from the mamillary body and the limbic cortex in the rat

Anterior thalamic afferents from the mamillary body and the limbic cortex were studied by using single and double retrograde transport methods in the rat. The medial mamillary nucleus was divided on the basis of the cytoarchitecture into four subnuclei: the pars medialis centralis, pars medialis dorsalis, pars lateralis, and pars basalis. Extensive connections were seen between each of these subdivisions of the mamillary body and the anterior thalamic nuclei, topographically organized so that the anteromedial thalamic nucleus receives projections exclusively from the pars medialis centralis, while the anteroventral thalamic nucleus receives projections from the pars medialis dorsalis and pars lateralis. Nuclei in the dorsal half of these two mamillary subdivisions project predominantly to the medial half of the anteroventral thalamic nucleus, and those in the ventral half to the lateral half of the nucleus. The pars basalis was found to have numerous projections to the magnocellular part of the anteroventral nucleus. All limbic cortical areas send projections bilaterally to all regions of the anteromedial nucleus as well as to the parvicellular parts of the anteroventral thalamic nucleus, while the anterodorsal nucleus receives ipsilateral projections originating exclusively from the preagranular, anterior limbic, and cingular regions. The magnocellular part of the anteroventral nucleus, however, receives only ipsilateral projections from all of the limbic cortex. Some neurons in the infralimbic region also project bilaterally to all of the anterior thalamic nuclei except the anterodorsal nucleus. All of these cortical projections to the anterior thalamus originate in layers V and VI of the limbic cortex.     

Thalamocortical projections of the anteroventral thalamic nucleus in the rabbit

The anterior thalamic nuclei are one of the regions that play critical roles in behavioral learning and memory functions. A part of the anterior thalamic nuclei, the anteroventral nucleus (AV) is well developed and differentiated into the parvocellular (AVp) and magnocellular (AVm) division in the rabbit. The AV is crucial for learning discriminative avoidance conditioning. Although communication between the AV and cortex is considered important in learning, little is known about the neural connections of the AV in the rabbit. Thus, this study used anterograde tracer biotinylated dextran amine and the retrograde tracer cholera toxin B subunit to examine the organization of the thalamocortical projections of the AV. Our data show that each division of the AV provides a unique set of projections to restricted regions and layers of the retrosplenial cortex and presubiculum. In addition, the AVp projects to layers I and IV of retrosplenial areas 29 and 30 and to layers I and VI of the presubiculum. The dorsolateral AVm projects to layers I and IV of area 29 and to layers I, III, and V of the presubiculum. However, the ventromedial AVm only projects to layer I of area 29. These projections are generally organized such that the rostral-to-caudal axis of the AV corresponds to the caudal-to-rostral axis of the retrosplenial cortex and to the temporal-to-septal axis of the presubiculum. These findings suggest distinct functional roles played by each division of the AV in the learning and memory functions. J. Comp. Neurol. 523:726–741, 2015. © 2014 Wiley Periodicals, Inc.     

Afferent connections of anterior thalamus in rats: sources and association with muscarinic acetylcholine receptors

Afferent connections of the anterior thalamic nuclei (ATN) are classically thought to originate in the mammillary body and limbic cortex. This study explores nonlimbic sources of ATN afferents by using retrograde transport of horseradish peroxidase (HRP) to ascertain the relative contribution of these connections. Spread of HRP into adjacent regions was prevented either by removing the overlying cortex or by injecting through permanently implanted cannulas. The main sources of nonlimbic ATN afferents are the pretectum and central gray. Pretectal neurons were HRP-labeled primarily in the contralateral medial pretectal nucleus with a smaller number in the ipsilateral posterior pretectal nucleus. In the central gray, labeled cells were concentrated ipsilaterally in the laterodorsal tegmental nucleus. Additional projections to ATN originate in the reticular and ventral lateral geniculate nuclei of the thalamus, raphe nuclei, peripontine tegmental nucleus, and locus coeruleus. The association of ATN afferents with muscarinic receptors was also explored by means of in vitro receptor autoradiography with the muscarinic ligands propylbenzilylcholine mustard (PrBCM) and pirenzepine (PZ) in normal rats and rats with ablations. Ibotenic acid injections into ATN were used to destroy intrinsic neurons while leaving afferent fibers intact. Whereas such ablations produced statistically significant decreases in PrBCM binding in the anterior dorsal (AD, -45%) and anterior ventral, magnocellular part (AVm, -51%) nuclei, binding in the anterior ventral, parvicellular part (AVp) and anterior medial (AM) nuclei was not significantly decreased. Furthermore, PZ binding in normal rat ATN was significantly less (-72%) than PrBCM binding. These results suggest that a major proportion of muscarinic binding is associated with presynaptic elements. Ibotenic acid ablations of the mammillary body reduced PrBCM binding in ATN whereas lesions in cingulate cortex and laterodorsal tegmental nucleus had no effect. Compared to sham lesion controls, mammillary body lesions resulted in statistically significant decreases in binding bilaterally in AD (-15%), AVm (-19%), and AM (-20%). In conclusion, ATN receive afferents from several nonlimbic regions. Of these inputs, the pretectum may be the primary route through which sensory information reaches ATN. In addition, cholinergic input may modulate activity in projections from the mammillary body to ATN through presynaptic muscarinic receptors.     

Organization of cortical afferents to the rostral,limbic sector of the rat thalamic reticular nucleus

The organization of limbic cortical afferents to the thalamic reticular nucleus (TRN) is described. Wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP), biocytin, neurobiotin, or fluorescent dextrans was delivered into the rat cingulate, retrosplenial, and, for comparison, somatosensory cortices, In other species a slab-like arrengement of cortical terminals has been described for sensory TRN sectors. Here this is seen in the rat somatosen-sory sector. Terminals from limbic cortices did not cluster into slabs but were found to fill the entire thickness of distinct rostral TRN regions. The cingulate and retrosplenial recipient TRN regions overlap, as do the projections from these cortical areas to anterior thalamic nuclei. Retrosplenial fibres contacted the dorsal and rostral TRN, which is be connected to the retrosplenial-recipient anteroventral, anterodorsal, and laterodorsal thalamic nucler. Cingulate terminals occupied more ventral regions of the rostral TRN. This area is connected to thalamic nuclei also innervated by the cingulate cortex: the mediodorsal and anteromedial nuclei. A loose, but clear, topography could be defined for the cingulate-reticular pathway: rostrocaudal and mediolateral directions in the cortex are represented by ventrodorsal and rostrocaudal directions in the TRN, respectively. This organization of limbic corticoreticular pathway corresponds to the arrangement of limbic corticothalamic connections. The ultrastruc-ture of the limbic cortical axon terminals was similar to that of the cortical boutons (D-type) described previously. The labelled terminals formed asymmetrical synapses onto dendritic profiles of reticular neurons. These findings, together with data in the literature, show significant morphological and connectional differences within the TRN that imply functional heterogeneities.     

Organization of projections of rat retrosplenial cortex to the anterior thalamic nuclei

The organization of the projections from the retrosplenial cortex (Brodmann's area 29) to the anterior thalamic nuclei was examined in the rat with retrograde transport of the cholera toxin B subunit and anterograde transport of biotinylated dextran amine. Areas 29a and 29b project mainly ipsilaterally to the rostral two‐thirds of the anteroventral nucleus, with area 29a projecting more rostrodorsally than area 29b. Area 29c projects bilaterally to the ventromedial part of the anteroventral nucleus. The projections from area 29c are organized in a topographic pattern such that the rostral area 29c projects to the caudoventral part of the anteroventral nucleus, whereas the caudal area 29c projects to the more rostrodorsal parts. Caudal area 29d projects mainly ipsilaterally to the rostrodorsal part of the anteromedial nucleus, and the rostral and dorsal parts of the anteroventral nucleus, whereas rostral area 29d projects bilaterally to the caudodorsal part of the anteromedial nucleus and the caudolateral part of the anteroventral nucleus. All the areas of the retrosplenial cortex provide sparse projections, mainly ipsilateral, to the anterodorsal nucleus, with a crude topographic pattern such that the rostrocaudal axis of the retrosplenial cortex corresponds to the caudorostral axis of the anterodorsal nucleus. The results indicate that each area of the retrosplenial cortex has a distinct projection field within the anterior thalamic nuclei. This suggests that each of these projections transmits distinct information that is important for complex memory and learning functions, e.g. discriminative avoidance learning and spatial memory.     

Efferent projections from the anterior thalamic nuclei to the cingulate cortex in the rat

The organization of projections from the anterior thalamic nuclei to the cingulate cortex was analyzed in the rat by the anterograde transport of Phaseolus vulgaris-leucoagglutinin. The rostral part of the anteromedial nucleus projects to layers I, V and VI of the anterior cingulate areas 1 and 2, layers I and III of the ventral orbital area, layers I, V and VI of area 29D of the retrosplenial area, and layers I and V of the caudal part of the retrosplenial granular and agranular areas. In contrast, the caudal part of the anteromedial nucleus projects to layer V of the frontal area 2, and layers I and V of the rostral part of the retrosplenial granular and agranular areas. The interanteromedial nucleus projects to layers I, III and V of the frontal area 2, layer V of the agranular insular area, and layers I, V and VI of area 29D. The anteroventral nucleus projects to layers I and IV of the retrosplenial granular area, whereas the anterodorsal nucleus projects to layers I, III and IV of the same area. Projections from the anteroventral and anterodorsal nuclei were, furthermore, organized such that their ventral parts project to the rostral part of the retrosplenial granular area, whereas their dorsal parts project to the more caudal part. The results suggest that the anterior thalamic nuclei project to more widespread areas and laminae of the cingulate cortex than was previously assumed. The projections are organized such that the anteromedial and interanteromedial nuclei project to layer I and the deep layers of the anterior cingulate and retrosplenial cortex, whereas the anteroventral and anterodorsal nuclei project to the superficial layers of the retrosplenial cortex. These thalamocortical projections may play important roles in behavioral learning such as discriminative avoidance behavior.     

Medial cortex of the lizard Gekko gecko: A hodological study with emphasis on regional specialization

There is increasing evidence that the archicortex in mammals and reptiles is not a homogeneous structure. However, little is known about the regional specialization of this cortical area in reptiles. Therefore, the efferent connections of the medial cortex of the lizard Gekko gecko were studied with the anterograde tracer Phaseolus vulgaris-leucoagglutinin. The small-celled part of the medial cortex (Cxms) projects to various parts of the septum in a topological way: the rostral part projects to the anterior septal nucleus, whereas the caudal part projects to the lateral septal nucleus and the nucleus septi impar. In addition, Cxms projects to the large-celled part of the medial cortex (Cxml). Axons that originate from the dorsal part of Cxms terminate at the proximal parts of the apical and basal dendrites of the neurons of Cxml caudal to the injection site. In contrast, fibers originating from the ventral part terminate on more distal parts of the dendrites of neurons of Cxml rostral to the injection site. Other projections from Cxms to the dorsal cortex (Cxd) and the external amygdaloid nucleus were found. The Cxml projects bilaterally to Cxms. These projections terminate in the superficial and deep plexiform layers. In addition, projections to the cell plate of Unger, Cxd, and to the lateral septal nucleus were found. It appears, on the basis of the efferent connections, that Cxms can be divided into a rostral and caudal part, while hodological differences also exist between the dorsal and ventral parts of Cxms. The results of the present study do not suggest a subdivision of Cxml. The regional variations of the medial cortex in the lizard Gekko gecko differ from the regional variations described in other reptilian species. © 1993 Wiley-Liss, Inc.     

Organization of thalamic projections to the ventral striatum in the primate

Although thalamic projections to the dorsal striatum are well described in primates and other species, little is known about thalamic projections to the ventral or “limbic” striatum in the primate. This study explores the organization of the thalamic projections to the ventral striatum in the primate brain by means of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) and Lucifer yellow (LY) retrograde tracer techniques. In addition, because functional and connective differences have been described for the core and shell components of the nucleus accumbens in the rat and are thought to be similar in the primate, this study also explores whether these regions of the nucleus accumbens can be distinguished by their thalamic input. Tracer injections are placed in different portions of the ventral striatum, including the medial and lateral regions of the ventral striatum; the central region of the ventral striatum, including the dorsal part of the core of the nucleus accumbens; and the shell region of the nucleus accumbens. Retrogradely labeled neurons are located mainly in the midline nuclear group (anterior and posterior paraventricular, paratenial, rhomboid, and reuniens thalamic nuclei) and in the parafascicular thalamic nucleus. Additional labeled cells are found in other portions of the intralaminar nuclear group as well as in other thalamic nuclei in the ventral, anterior, medial, lateral, and posterior thalamic nuclear groups. The distribution of labeled cells varies depending on the area of the ventral striatum injected. All regions of the ventral striatum receive strong projections from the midline thalamic nuclei and from the parafascicular nucleus. In addition, the medial region of the ventral striatum receives numerous projections from the central superior lateral nucleus, the magnocellular subdivision of the ventral anterior nucleus, and parts of the mediodorsal nucleus. After injection into the lateral region of the ventral striatum, few labeled neurons are seen scattered in nuclei of the intralaminar and ventral thalamic groups and occasional labeled cells in the mediodorsal nucleus. The central region of the ventral striatum, including the dorsal part of the core of the nucleus accumbens, receives a limited projection from the midline thqlamic, predominantly from the rhomboid nucleus. It receives much smaller projections from the central medial nucleus and the ventral, anterior, and medial thalamic groups. The shell of the nucleus accumbens receives the most limited projection from the thalamus and is innervated almost exclusively by the midline thalamic nuclei and the central medial and parafascicular nuclei. The shell is distinguished from the rest of the ventral striatum in that it receives the fewest projections from the ventral, anterior, medial, and lateral thalamic nuclei. © 1995 Wiley-Liss, Inc.     

Prosencephalic afferents to the mediodorsal thalamic nucleus

The afferent projections from the prosencephalon to the mediodorsal thalamic nucleus (MD) were studied in the cat by use of the method of retrograde transport of horseradish peroxidase (HRP). Cortical and subcortical prosencephalic structures project bilaterally to the MD. The cortical afferents originate mainly in the ipsilateral prefrontal cortex. The premotor, prelimbic, anterior limbic, and insular agranular cortical areas are also origins of consistent projections to the MD. The motor cortex, insular granular area, and some other cortical association areas may be the source of cortical connections to the MD. The subcortical projections originate principally in the ipsilateral rostral part of the reticular thalamic nucleus and the rostral lateral hypothalamic area. Other parts of the hypothalamus, the most caudal parts of the thalamic reticular nucleus, the basal prosencephalic structures, the zona incerta, the claustrum, and the entopeduncular and subthalamic nuclei are also sources of projections to the MD. Distinct, but somewhat overlapping areas of the prosencephalon project to the three vertical subdivisions of MD (medial, intermediate, and lateral). The medial band of the MD receives a small number of prosencephalic projections; these arise mainly in the caudal and ventral parts of the prefrontal cortex. Cortical projections also arise in the infralimbic area, while subcortical projections originate in the medial part of the rostral reticular thalamic nucleus and lateral hypothalamic area. The intermediate band of the MD receives the largest number of fibers from the prosencephalon. These arise principally in the intermediate and dorsal part of the lateral and medial surface of the prefrontal cortex, the premotor cortex, and the prelimbic and agranular insular areas. Projections also originate in basal prosencephalic formations (preoptic area, Broca's diagonal band, substantia innominata, and olfactory tubercle), rostral reticular thalamic nucleus, and lateral hypothalamic area. A large number of prosencephalic structures also project to the lateral band of the MD. These are mainly the most dorsal and caudal parts of the lateral and medial surface of the prefrontal cortex, the premotor and motor cortices, and the prelimbic, anterior limbic, and insular areas. Projections arise also in the lateral rostral and caudal parts of the reticular thalamic nucleus, the zona incerta, the lateral and dorsal hypothalamic areas, the claustrum, and the entopeduncular nucleus. These and previous results demonstrate a gradation in the afferent connections to the three subdivisions of the MD. Brain structures related to the olfactory sensory modality and with allocortical formations of the limbic system project principally to the medial band of the MD. The intermediate band of the MD receives subcortical and cortical projections from structures mainly related to the limbic system and cortical regions related to sensory association cortices. The lateral band of the MD receives projections mainly originating in structures related to complex sensory associative processes and to the motor system (especially from brainstem and cortical structures implicated in the regulation of eye movements).     

Retrograde double-labeling study of the mammillothalamic and the mammillotegmental projections in the rat

Collateral axonal branching from the medial or lateral mammillary nuclei to the anterior thalamus, Gudden's tegmental nuclei, the nucleus reticularis tegmenti pontis, and the medial pontine nucleus was studied using the fluorescent retrograde double-labeling method. One day after injection of Fast Blue into the anterior thalamic nuclei or Gudden's tegmental nuclei, Nuclear Yellow was injected into Gudden's tegmental nuclei or the nucleus reticularis tegmenti pontis and the medial pontine nucleus. Following 1 day survival, single- and double-labeled neurons were examined in the mammillary nuclei. The lateral mammillary nucleus contains neurons whose collateral fibers project to both the dorsal tegmental nucleus of Gudden and the ipsilateral or contralateral anterodorsal thalamic nucleus, to both the medial pontine nucleus and the anterodorsal thalamic nucleus, and to both the dorsal tegmental nucleus of Gudden and the medial pontine nucleus. The pars medianus and pars medialis of the medial mammillary nucleus contain neurons whose collateral fibers project to both the anteromedial thalamic nucleus and the ventral tegmental nucleus of Gudden, to both the anteromedial thalamic nucleus and the medial part of the nucleus reticularis tegmenti pontis, and to both the ventral tegmental nucleus of Gudden and the medial part of the nucleus reticularis tegmenti pontis. The dorsal half of the pars posterior of the medial mammillary nucleus contains a few neurons whose collateral fibers project to both the anteromedial thalamic nucleus and the rostral part of the ventral tegmental nucleus of Gudden, and to both the caudal part of the anteroventral thalamic nucleus and the rostral part of the ventral tegmental nucleus of Gudden, while the pars lateralis of the medial mammillary nucleus contains no double-labeled neurons and projects only to the anteroventral thalamic nucleus.     

Afferent projections to the interpeduncular nucleus in the rat, as studied by retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase

We examined the afferent projections to the subnuclei of the interpeduncular nucleus (IPN) in the rat by means of retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). We observed locations of retrogradely labeled cells following injections of WGA-HRP into the IPN, and distributions of anterogradely labeled fibers and terminals within the IPN following injections into the areas that contain cells of origin of afferents. Results of the retrograde and anterograde experiments have clarified the detailed organization of the IPN afferents. A part of the nucleus incertus, located dorsomedial to the dorsal tegmental nucleus, projects to the contralateral half of the rostral subnucleus of the IPN; the pars caudalis of the dorsal tegmental nucleus projects sparsely to the rostral lateral, dorsal lateral, lateral, caudal, and apical subnuclei predominantly contralaterally; the laterodorsal tegmental nucleus, to most of the subnuclei predominantly contralaterally; the ventromedial central gray rostral to the dorsal tegmental nucleus and lateral to the dorsal raphe nucleus projects to the rostral lateral and dorsal lateral subnuclei predominantly contralaterally; the median raphe nucleus, substantially to all subnuclei; the medial habenular nucleus, in a topographic manner, to the rostral, central, and intermediate subnuclei, to the rostral lateral and lateral subnuclei predominantly ipsilaterally, and to the dorsal lateral subnucleus predominantly contralaterally; the supramammillary nucleus and areas around the origin of the mammillothalamic tract and near the third ventricle project sparsely to the ventral part of the rostral subnucleus and to the central, lateral, caudal and apical subnuclei; the nucleus of the diagonal band, sparsely to the rostral, central, dorsal lateral, caudal, and apical subnuclei. These differential projections of the afferents to the subnuclei of the IPN may reflect its complex functions within the limbic midbrain circuit.     

Connections of the parabrachial nucleus with the nucleus of the solitary tract and the medullary reticular formation in the rat.

We examined the subnuclear organization of projections to the parabrachial nucleus (PB) from the nucleus of the solitary tract (NTS), area postrema, and medullary reticular formation in the rat by using the anterograde and retrograde transport of wheat germ agglutinin-horseradish peroxidase conjugate and anterograde tracing with Phaseolus vulgaris-leucoagglutinin. Different functional regions of the NTS/area postrema complex and medullary reticular formation were found to innervate largely nonoverlapping zones in the PB. The general visceral part of the NTS, including the medial, parvicellular, intermediate, and commissural NTS subnuclei and the core of the area postrema, projects to restricted terminal zones in the inner portion of the external lateral PB, the central and dorsal lateral PB subnuclei, and the "waist" area. The dorsomedial NTS subnucleus and the rim of the area postrema specifically innervate the outer portion of the external lateral PB subnucleus. In addition, the medial NTS innervates the caudal lateral part of the external medial PB subnucleus. The respiratory part of the NTS, comprising the ventrolateral, intermediate, and caudal commissural subnuclei, is reciprocally connected with the K?lliker-Fuse nucleus, and with the far lateral parts of the dorsal and central lateral PB subnuclei. There is also a patchy projection to the caudal lateral part of the external medial PB subnucleus from the ventrolateral NTS. The rostral, gustatory part of the NTS projects mainly to the caudal medial parts of the PB complex, including the "waist" area, as well as more rostrally to parts of the medial, external medial, ventral, and central lateral PB subnuclei. The connections of different portions of the medullary reticular formation with the PB complex reflect the same patterns of organization, but are reciprocal. The periambiguus region is reciprocally connected with the same PB subnuclei as the ventrolateral NTS; the rostral ventrolateral reticular nucleus with the same PB subnuclei as both the ventrolateral (respiratory) and medial (general visceral) NTS; and the parvicellular reticular area, adjacent to the rostral NTS, with parts of the central and ventral lateral and the medial PB subnuclei that also receive rostral (gustatory) NTS input. In addition, the rostral ventrolateral reticular nucleus and the parvicellular reticular formation have more extensive connections with parts of the rostral PB and the subjacent reticular formation that receive little if any NTS input. The PB contains a series of topographically complex terminal domains reflecting the functional organization of its afferent sources in the NTS and medullary reticular formation.     

Organization in the somatosensory sector of the cat's thalamic reticular nucleus

This study describes the organization of cells in the thalamic reticular nucleus (TRN) that project to the somatosensory part of the dorsal thalamus in the cat. Injections of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) and fluorescent dyes were made into the ventrobasal complex (VB) and the medial division of the posterior complex (POm) of the thalamus. The resultant retrograde labelling in TRN was analyzed. Large injections of a tracer in VB label many reticular cells that are restricted to a centroventral, or somatosensory, sector of TRN. Small injections of a tracer in VB produce narrow zones of labelled cells in this sector. In reconstructions these zones resemble thin “slabs,” which lie parallel to the plane of TRN along its oblique rostrocaudal dimension and occupy only a fraction of its thickness. In comparisons of the zones of labelled cells in TRN resulting from tracer injections in different nuclei of VB, inner cells, intermediate cells, and outer cells across the thickness of TRN project to the ventral posteromedial, the medial division of the ventral posterolateral, and the lateral division of the ventral posterolateral nuclei, respectively. Furthermore, shifts in injected areas along the dorsoventral dimension of VB produce similar shifts in zones of labelled cells in TRN. Thus, reticular cells form an accurate map on the basis of their connections with VB. Large injections of a tracer in the ventral subdivision of POm label many reticular cells that are also restricted to the centroventral sector of TRN. Small injections of a tracer in ventral POm produce broad zones of labelled cells in this sector. In comparisons of the zones of labelled cells in TRN resulting from tracer injections in different regions of ventral POm, cells that project to these regions are scattered across the thickness of TRN and occupy overlapping territories. Large injections of a tracer in either VB or ventral POm also label cells in a restricted centroventral region of the perireticular nucleus. Double injections of different tracers in VB and ventral POm produce many cells in TRN that are labelled from both of these dorsal thalamic structures and fewer cells that are labelled from only one or the other of these structures. These results indicate that there is a dual organization in the projections of cells in the somatosensory sector of TRN to dorsal thalamus: Projections to VB are topographically organized whereas those to ventral POm lack a topographical organization. Furthermore, both of these mapped and nonmapped projections can arise from single reticular cells in the somatosensory sector. © 1996 Wiley-Liss, Inc.     

Brainstem projections to midline and intralaminar thalamic nuclei of the rat

The projections from the brainstem to the midline and intralaminar thalamic nuclei were examined in the rat. Stereotaxic injections of the retrograde tracer cholera toxin beta -subunit (CTb) were made in each of the intralaminar nuclei of the dorsal thalamus: the lateral parafascicular, medial parafascicular, central lateral, paracentral, oval paracentral, and central medial nuclei; in the midline thalamic nuclei-the paraventricular, intermediodorsal, mediodorsal, paratenial, rhomboid, reuniens, and submedius nuclei; and, in the anteroventral, parvicellular part of the ventral posterior, and caudal ventral medial nuclei. The retrograde cell body labeling pattern within the brainstem nuclei was then analyzed. Nearly every thalamic site received a projection from the deep mesencephalic reticular, pedunculopontine tegmental, dorsal raphe, median raphe, laterodorsal tegmental, and locus coeruleus nuclei. Most intralaminar thalamic sites were also innervated by unique combinations of medullary and pontine reticular formation nuclei such as the subnucleus reticularis dorsalis, gigantocellular, dorsal paragigantocellular, lateral, parvicellular, caudal pontine, ventral pontine, and oral pontine reticular nuclei; the dorsomedial tegmental, subpeduncular tegmental, and ventral tegmental areas; and, the central tegmental field. In addition, most intralaminar injections resulted in retrograde cell body labeling in the substantia nigra, nucleus Darkschewitsch, interstitial nucleus of Cajal, and cuneiform nucleus. Details concerning the pathways from the spinal trigeminal, nucleus tractus solitarius, raphe magnus, raphe pallidus, and the rostral and caudal linear raphe nuclei to subsets of midline and intralaminar thalamic sites are discussed in the text. The discussion focuses on brainstem-thalamic pathways that are likely involved in arousal, somatosensory, and visceral functions.     

Organization of the pallium in the fire-bellied toad Bombina orientalis. I: Morphology and axonal projection pattern of neurons revealed by intracellular biocytin labeling

The cytoarchitecture and axonal projection pattern of pallial areas was studied in the fire-bellied toad Bombina orientalis by intracellular injection of biocytin into a total of 326 neurons forming 204 clusters. Five pallial regions were identified, differing in morphology and projection pattern of neurons. The rostral pallium receiving the bulk of dorsal thalamic afferents has reciprocal connections with all other pallial areas and projects to the septum, nucleus accumbens, and anterior dorsal striatum. The medial pallium projects bilaterally to the medial pallium, septum, nucleus accumbens, mediocentral amygdala, and hypothalamus and ipsilaterally to the rostral, dorsal, and lateral pallium. The ventral part of the medial pallium is distinguished by efferents to the eminentia thalami and the absence of contralateral projections. The dorsal pallium has only ipsilateral projections running to the rostral, medial, and lateral pallium; septum; nucleus accumbens; and eminentia thalami. The lateral pallium has ipsilateral projections to the olfactory bulbs and to the rostral, medial, dorsal, and ventral pallium. The ventral pallium including the striatopallial transition area (SPTA) has ipsilateral projections to the olfactory bulbs, rostral and lateral pallium, dorsal striatopallidum, vomeronasal amygdala, and hypothalamus. The medial pallium can be tentatively homologized with the mammalian hippocampal formation, the dorsal pallium with allocortical areas, the lateral pallium rostrally with the piriform and caudally with the entorhinal cortex, the ventral pallium with the accessory olfactory amygdala. The rostral pallium, with its projections to the dorsal and ventral striatopallidum, resembles the mammalian frontal cortex.     

Topographic organization of the projections from the interstitial system of the spinal trigeminal tract to the parabrachial nucleus in the rat

Neurons in the paratrigeminal nucleus are known to project to the parabrachial region, but both these areas are heterogeneous, and the subnuclei that account for these connections are not known. To characterize better these projections, we injected small amounts of fluorogold or latex beads labeled with rhodamine or fluorescein into the parabrachial area in the rat and evaluated the retrograde transport of tracer to the paratrigeminal nucleus and neighboring regions. The results show that the rostral part of the paratrigeminal nucleus projects to the medial subnucleus of the parabrachial nucleus. The intermediary part of the paratrigeminal nucleus projects to both the external lateral and to the external medial subnuclei of the parabrachial nucleus. The caudal part of the paratrigeminal nucleus projects to the ventral lateral subnucleus of the parabrachial nucleus. The dorsal paramarginal nucleus projects to the external lateral and the extreme lateral subnuclei of the parabrachial nucleus. Lamina I and II of the spinal trigeminal nucleus also project to the external lateral and the extreme lateral subnuclei of the parabrachial nucleus. In conclusion, the rostral, intermediate, and caudal parts of the paratrigeminal nucleus and the dorsal paramarginal nucleus each have clearly different projection patterns and presumably have different functions.     

The cortical projections of the mediodorsal nucleus and adjacent thalamic nuclei in the rat.

The mediodorsal nucleus of the rat thalamus has been divided into medial, central and lateral segments on the basis of its structure and axonal connections, and these segments have been shown by experiments using the autoradiographic method of demonstrating axonal connections to project to seven distinct cortical areas covering most of the frontal pole of the hemisphere. The position and cytoarchitectonic characteristics of these areas are described. The medial segment of the nucleus projects to the prelimbic area (32) on the medial surface of the hemisphere, and to the dorsal agranular insular area, dorsal to the rhinal sulcus on the lateral surface. The lateral segment projects to the anterior cingulate area (area 24) and the medial precentral area on the dorsomedial shoulder of the hemisphere, while the central segment projects to the ventral agranular insular area in the dorsal bank of the rhinal sulcus, and to a lateral part of the orbital cortex further rostrally. (The term "orbital" is used to refer to the cortex on the ventral surface of the frontal pole of the hemisphere.) A ventral part of this orbital cortex also receives fibers from the mediodorsal nucleus, possibly its lateral segment, but the medial part of the orbital cortex, and the ventrolateral orbital area in the fundus of the rhinal sulcus receive projections from the paratenial nucleus and the submedial nucleus, respectively. All of these thalamocortical projections end in layer III, and in the outer part of layer I. The basal nucleus of the ventromedial complex (the thalamic taste relay) has been shown to have a similar laminar projection (layer I and layers III/IV) to the granular insular area immediately dorsal to, but not overlapping, the mediodorsal projection field. However, the principal nucleus of the ventromedial complex appears to project to layer I, and possibly layer VI, of the entire frontal pole of the hemisphere. The anteromedial nucleus does not appear to project to layer III of the projection field of the mediodorsal nucleus, although it may project to layers I and VI, especially in the anterior cingulate and medial precentral areas. A thalamoamygdaloid projection from the medial segment of the mediodorsal nucleus to the basolateral nucleus of the amygdala has also been demonstrated, which reciprocates an amygdalothalamic projection from the basolateral nucleus to the medial segment. The habenular nuclei also appear to project to the central nucleus of the amygdala. These results are discussed in relation to the delineation and subdivision of the prefrontal cortex in the rat, and to amygdalothalamic and amygdalocortical projections which are described in a subsequent paper (Krettek and Price, '77).