Corticothalamic connections from the second somatosensory area and neighboring regions in the lateral sulcus of macaque monkeys

Corticothalamic connections were shown between the second somatosensory area in primates and the ventroposterior nuclei of the thalamus. These projections were topographically arranged with those from the hindlimb portions of SII traced to the most lateral and posterior parts of the ventroposterior lateral nucleus (VPLc) and those from the forelimb located medially within VPLc. The densest labeling was found ventrally in VPLc and dorsally within ventroposterior inferior n. (VPI) only after injections of the forelimb. A more scattered, dorsal distribution of labeling was seen in the rest of VPLc from injections involving more proximal parts of the body representation in SII.     

Connections between area 3b of the somatosensory cortex and subdivisions of the ventroposterior nuclear complex and the anterior pulvinar nucleus in squirrel monkeys.

The goal of this study was to determine whether somatosensory thalamic nuclei other than the ventroposterior nucleus proper (VP) have connections with area 3b of the postcentral cortex in squirrel monkeys. Small injections of the anatomical tracers wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) or 3H-proline were placed in electrophysiologically identified representations of body parts. The results indicate that, besides the well-established somatotopically organized connections with VP, area 3b has connections with three other nuclei of the somatosensory thalamus: the ventroposterior superior nucleus (VPS ["shell" of VP]), the ventroposterior inferior nucleus (VPI), and the anterior pulvinar nucleus (Pa). Injections confined to area 3b or involving adjacent parts of area 3a or area 1 indicate that connections between VPS, VPI, and Pa and the postcentral cortex are somatotopically organized. In VPS, connections related to the hand were found medially, and connections related to the foot were lateral. In VPI, connections with the cortical representations of the mouth, hand, and foot were successively more lateral. In Pa, connections related to the mouth, hand, and foot were successively more ventral, lateral, and caudal, and the trunk region was caudomedial. The findings suggest that VPI contains a representation of all parts of the body, including the face. The connections of Pa with the primary somatosensory cortex, area 3b, the location of Pa relative to the ventroposterior nucleus, and the high degree of topographic order in the connections of Pa with the postcentral cortex suggest that Pa is an integral part of the somatosensory thalamus in monkeys and is homologous to the medial nucleus of the posterior group (Pom) in other mammals. Overall, the results contribute to the growing evidence that individual somatosensory cortical areas in monkeys receive inputs from multiple thalamic sources, and that a single thalamic nucleus has several cortical targets.     

Thalamic connections of three representations of the body surface in somatosensory cortex of gray squirrels

The anatomical tracer, wheat germ agglutinin, was used to determine the connections of electrophysiologically identified locations in three architectonically distinct representations of the body surface in the somatosensory cortex of gray squirrels. Injections in the first somatosensory area, S-I, revealed reciprocal connections with the ventroposterior nucleus (VP), a portion of the thalamus just dorsomedial to VP, the posterior medial nucleus, Pom, and sometimes the ventroposterior inferior nucleus (VPI). As expected, injections in the representation of the face in S-I resulted in label in ventroposterior medial (VPM), the medial subnucleus of VP, whereas injections in the representation of the body labeled ventroposterior lateral (VPL), the lateral subnucleus of VP. Furthermore, there was evidence from connections that the caudal face and head are represented dorsolaterally in VPM, and the forelimb is represented centrally and medially in VPL. The results also support the conclusion that a representation paralleling that in VP exists in Pom, so that the ventrolateral part of Pom represents the face and the dorsomedial part of Pom is devoted to the body. Because connections with VPI were not consistently revealed, the possibility exists that only some parts or functional modules of S-I are interconnected with VPI. Two separate small representations of the body surface adjoin the caudoventral border of S-I. Both resemble the second somatosensory area, S-II, enough to be identified as S-II in the absence of evidence for the other. We term the more dorsal of the two fields S-II because it was previously defined as S-II in squirrels (Nelson et al., '79), and because it more closely resembles the S-II identified in most other mammals. We refer to the other field as the parietal ventral area, PV (Krubitzer et al, '86). Injections in S-II revealed reciprocal connections with VP, Pom, and a thalamic region lateral and caudal to Pom and dorsal to VP, the posterior lateral nucleus, Pol. Whereas major interconnections between S-II and VPI have been reported for cats, raccoons, and monkeys, no such interconnections were found for S-II in squirrels. The parietal ventral area, PV, was found to have prominent reciprocal interconnections with VP, VPI, and the internal (magnocellular) division of the medial geniculate complex (MGi). The pattern of connections conforms to the established somatotopic organization of VP and suggests a crude parallel somatotopic organization in VPI. Less prominent interconnections were with Pol.(ABSTRACT TRUNCATED AT 400 WORDS)     

Thalamic Input to Representations of the Teeth,Tongue, and Face in Somatosensory Area 3b of Macaque Monkeys

Representations of the parts of the oral cavity and face in somatosensory area 3b of macaque monkeys were identified with microelectrode recordings and injected with different neuroanatomical tracers to reveal patterns of thalamic projections to tongue, teeth, and other representations in primary somatosensory cortex. The locations of injection sites and resulting labeled neurons were further determined by relating sections processed to reveal tracers to those processed for myeloarchitecture in the cortex and multiple architectural stains in the thalamus. The ventroposterior medial subnucleus (VPM) for touch was identified as separate from the ventroposterior medial parvicellular nucleus (VPMpc) for taste by differential expression of several types of proteins. Our results revealed somatotopically matched projections from VPM to the part of 3b representing intra‐oral structures and the face. Retrogradely labeled cells resulting from injections in area 3b were also found in other thalamic nuclei including: anterior pulvinar (Pa), ventroposterior inferior (VPI), ventroposterior superior (VPS), ventroposterior lateral (VPL), ventral lateral (VL), center median (CM), central lateral (CL), and medial dorsal (MD). None of our injections, including those into the representation of the tongue, labeled neurons in VPMpc, the thalamic taste nucleus. Thus, area 3b does not appear to be involved in processing taste information from the thalamus. This result stands in contrast to those reported for New World monkeys. J. Comp. Neurol. 521:3954–3971, 2013. © 2013 Wiley Periodicals, Inc.     

Multisensory convergence in auditory cortex, II. Thalamocortical connections of the caudal superior temporal plane

Recent studies of macaque monkey auditory cortex have revealed convergent auditory and somatosensory activity in the caudomedial area (CM) of the belt region. In the present study and its companion (Smiley et al., J. Comp. Neurol. [this issue]), neuroanatomical tracers were injected into CM and adjacent areas of the superior temporal plane to identify sources of auditory and somatosensory input to this region. Other than CM, target areas included: A1, caudolateral belt (CL), retroinsular (Ri), and temporal parietotemporal (Tpt). Cells labeled by injections of these areas were distributed mainly among the ventral (MGv), posterodorsal (MGpd), anterodorsal (MGad), and magnocellular (MGm) divisions of the medial geniculate complex (MGC) and several nuclei with established multisensory features: posterior (Po), suprageniculate (Sg), limitans (Lim), and medial pulvinar (PM). The principal inputs of CM were MGad, MGv, and MGm, with secondary inputs from multisensory nuclei. The main inputs of CL were Po and MGpd, with secondary inputs from MGad, MGm, and multisensory nuclei. A1 was dominated by inputs from MGv and MGad, with light multisensory inputs. The input profile of Tpt closely resembled that of CL, but with reduced MGC inputs. Injections of Ri also involved CM but strongly favored MGm and multisensory nuclei, with secondary inputs from MGC and the inferior division (VPI) of the ventroposterior complex (VP). The results indicate that the thalamic inputs of areas in the caudal superior temporal plane arise mainly from the same nuclei, but in different proportions. Somatosensory inputs may reach CM and CL through MGm or the multisensory nuclei but not VP.     

Spinothalamocortical projections to the secondary somatosensory cortex (SII) in squirrel monkey

Anterograde labeling of the cervical spinothalamic tract was combined with retrograde labeling of thalamocortical cells projecting to the hand region of the second somatosensory cortex (hSII) to identify likely sites in the thalamus for processing and transmitting nociceptive information to hSII. Anterograde labeling of terminals was done with 2% WGA-HRP injections in the cervical enlargement; thalamocortical cells were retrogradely labeled with fluorescent tracers. In one experiment, the contralateral primary somatosensory cortex hand region (hSI) was injected to provide a direct comparison with hSII thalamic label. Both labeled cells and terminal-like structures were visualized in single thalamic sections and their numbers and positions quantitatively analyzed. The number of labeled cells within 100 microns from the STT terminals were counted as overlapping cells. Four thalamic nuclei, ventroposterior inferior (VPI), ventroposterior lateral (VPL), posterior nucleus (PO) and centrolateral nucleus (CL) combined to contain 86.5% of all hSII-projecting overlapping cells. Of all hSII-projecting thalamic overlapping cells, VPI contained the largest number (36.4% of the total) followed by the anterior portion of the posterior nuclear complex (POa; 20.4%), VPL (18.3%) and CL (11.4%). Results of the hSI injection show a different pattern of overlap in agreement with our earlier study. The relative distribution of overlapping cells was dependent on the antero-posterior position of the SII injections. The most anterior injections resulted in small numbers of labeled cells, with the majority of overlapping cells located in PO and CL. The more posterior injections resulted in overlapping cells mainly in VPI and VPL. The results indicate that, in the squirrel monkey, VPI, VPL, POa and CL relay nociceptive information from the spinal cord to the second somatosensory cortex.     

Thalamic projections to fields A, AI, P, and VP in the cat auditory cortex

Thalamocortical projections to four tonotopic fields (A, AI, P, and VP) of the cat auditory cortex were studied by using combined microelectrode mapping and retrograde axonal transport techniques. Horseradish peroxidase (HRP) or HRP combined with either tritiated bovine serum albumin or nuclear yellow was injected into identified best-frequency sites of one or two different fields in the same brain. Arrays of labeled neurons were related to thalamic nuclei defined on the basis of their cytoarchitecture and physiology. In some cases, patterns of labeling were directly compared with thalamic best-frequency maps obtained in the same brain. We compared only patterns of labeling resulting from injections into similar parts of the frequency representation in different fields to insure that observed differences in patterns of labeling did not simply reflect differences in the frequency representation at the injection sites. The thalamic projection to the four fields is divided among seven nuclei, three tonotopic nuclei (ventral nucleus, V; lateral part of the posterior group of thalamic nuclei, Po; and dorsal cap nucleus, d) and four nontonotopic nuclei (caudodorsal nucleus, cd; ventrolateral nucleus, vl; and small, Ms; and medium-large, Mg, cell regions of the medial division). Projections to each field differ, and each field receives inputs from tonotopic and nontonotopic nuclei. Field A receives its major inputs from Po and Mg, and a minor input from V. Field AI receives its major inputs from V, Po, and Mg, although Po and Mg have heavier projections to field A. Field P receives its major inputs from V, d, and vl; and minor inputs from cd, Ms, Mg, and Po. Field VP receives major inputs from V, vl, and cd; and minor inputs from d, Ms, and Mg. There are segregated territories in V and Po in which most neurons projects to one cortical field (major projection), and a smaller proportion projects to one or more other fields (minor projections). Field VP receives a major projection from the caudal pole of V. Field P receives a major projection from the caudal half of V, and from a thin band along the dorsal border of rostral V. Field AI receives a major projection from most of the rostral one-half of V, and smaller areas in Po and the caudal half of V exclusive of its caudal pole. Field A receives a major projection from most of Po.(ABSTRACT TRUNCATED AT 400 WORDS)     

Cortical and thalamic connections of the parietal ventral somatosensory area in marmoset monkeys (Callithrix jacchus).

Microelectrode mapping methods were used to define the parietal ventral somatosensory area (PV) on the upper bank of the lateral sulcus in five marmosets (Callithrix jacchus). In the same animals, neuroanatomical tracers were placed into electrophysiologically identified sites in PV and/or the second somatosensory area (S2). Foci of anterograde and retrograde label were related to electrophysiological maps of cortical areas and cortical and thalamic architecture. The results lead to the following conclusions: (1) Multiunit recordings from cortex on the upper bank of the lateral sulcus demonstrate that PV is somatotopically organized, with the face representation adjoining area 3b and the hindlimb and tail representations away from this border in cortex deep on the upper bank of the lateral sulcus. The forelimb representation is caudal in PV adjacent to the S2 forelimb representation. The body surface representation in PV approximates a mirror image of that in S2; (2) Areas PV and S2 are less myelinated and have less cytochrome oxidase enzyme activity than area 3b; (3) The ventroposterior inferior nucleus (VPI) of the thalamus provides the major somatosensory projections to PV. PV is reciprocally connected with VPI and anterior pulvinar; (4) PV has ipsilateral cortical connections with areas 3a, 3b, 1, and M1 and higher order somatosensory fields, and at least most of these connections are somatotopically matched; and (5) Callosal connections of PV are with S2 and PV of the other cerebral hemisphere. These results further establish PV as one of at least four somatosensory areas of the lateral sulcus of primates.     

Thalamic connections of the vestibular cortical fields in the squirrel monkey (Saimiri sciureus).

The afferent thalamic connections to cortical fields important for control of head movement in space were analysed by intracortical retrograde tracer injections. The proprioceptive/vestibular area 3aV, the neck-trunk region of area 3a, receives two thirds of its thalamic projections from the oral and superior ventroposterior nucleus (VPO/VPS), which is considered as the proprioceptive relay of the ventroposterior complex (Kaas et al., J. Comp. Neurol. 226:211-240, 1984). The parieto-insular vestibular cortex (PIVC, area retroinsularis, Ri) receives its main thalamic input from posterior parts of the ventroposterior complex and from the medial pulvinar. Anatomical evidence is presented that the posterior region of the ventroposterior complex is a special compartment within this principal somatosensory relay complex. The parietotemporal association area T3, mainly involved in visual-optokinetic signal processing, receives a substantial input from the medial, the lateral, and the inferior pulvinar. Dual tracer experiments revealed that about 5% of the thalamic neurons projecting to 3aV were spatially intermingled with neurons projecting to areas PIVC or T3. This spatial intermingling was distributed over small but numerous, circumscribed thalamic regions, called "common patches," which were found mainly in the intralaminar nuclei, the posterior group of thalamic nuclei, and the caudal parts of the ventroposterior complex. The "common patches" may indicate a functional coupling of area 3aV with the PIVC or area T3 on the thalamic level. In control experiments thalamic projections to the granular insula Ig and the anterior part of area 7, two cerebral structures connected with the vestibular cortical areas, were studied. Some overlap in the thalamic relay structures projecting to these areas with those projecting to the vestibular cortices was found. A quantitative evaluation of thalamic regions projecting to different cortical structures was performed by constructing so-called "thalamograms." A scheme was developed that describes the afferent thalamic connections by which vestibular, visual-optokinetic, and proprioceptive signals reach the vestibular cortical areas PIVC and 3aV.     

The Somatotopic Organization Within the Cat's Thalamic Reticular Nucleus

The organization of the somatosensory representation within the cat's thalamic reticular nucleus (TRN) was studied. Focal injections of horseradish peroxidase (HRP), wheatgerm agglutinin conjugated to HRP, and/or [3H]proline were made into somatosensory cortical areas 1 (S1) and 2 (S2). The resultant labelling in the thalamus was analysed. Single injections into S1 result in single zones of terminal labelling in TRN that are restricted to the centroventral part of the sheet-like nucleus. In reconstructions from horizontal sections these zones of labelling resemble thin 'slabs', which lie in the plane of the nucleus parallel to its borders, occupy only a fraction of the thickness of the reticular sheet, and are broadly elongated in the dorsoventral and oblique rostrocaudal dimensions. Thus, the slabs of S1 terminals, which represent large loci of the body surface, and the main distribution of the reticular dendrites have a similar orientation. In comparisons of the zones of labelling following single or double injections at different cortical sites in S1, an inner (medial) to outer (lateral) shift in labelling in the ventrobasal complex (VB) is accompanied by an inner (medial) to outer (lateral) shift in labelling along the thickness of the reticular sheet. Thus, like VB the reticular nucleus receives a topographically accurate projection from S1. Further, the somatotopic map conveyed from S1 to TRN is orientated perpendicular to the plane of the nucleus and repeats the spatial organization of the map in VB. S2 injections result in zones of terminal labelling in that part of TRN that receives S1 inputs. On the basis of these findings, together with those in other mammalian species, two conclusions can be reached about corticoreticular relations. First, although there can be continuity in individual maps of cortical inputs to TRN, there are discontinuities in cortical representations at the inner and outer borders of the reticular sheet. Second, TRN can receive a significant convergence of inputs from different cortical areas.     

Connections of area 2 of somatosensory cortex with the anterior pulvinar and subdivisions of the ventroposterior complex in macaque monkeys

The principal goal of the present study was to determine the thalamic connections of area 2 of postcentral somatosensory cortex of monkeys. The placement of injections of anatomical tracers (horseradish peroxidase, wheat germ agglutinin, or ³H-proline) was guided by extensive microelectrode maps of cortex in the region of the injection site. These maps identified the body parts represented in the cortex included in the injection site, and provided information about the physiological boundaries of area 2, which was related later to the cortical architecture. Most injections were placed in the representation of the hand in area 2, which was highly responsive to cutaneous stimuli and could be mapped in detail. Injections were also placed in other parts of area 2, area 1, or area 5, and some injections involved more than one area. As other investigators have determined, regions of retrograde and anterograde thalamic label overlapped, demonstrating that connections with cortex are reciprocal. Injections completely confined to area 2 consistently produced label in two locations: the anterior pulvinar (Pa) and a dorsal capping zone of the ventroposterior complex that we term the ventroposterior superior nucleus (VPS). Single restricted injection sites resulted in one region of label in VPS, and multiple foci of label in Pa. In some cases where the injection was confined to the representation of the hand in area 2, label was also found more ventrally in the ventroposterior complex in ventroposterior nucleus proper (VP). Thus, area 2 receives input from Pa, VPS, and, at least in some locations and individuals, VP. Injections of tracers into area 1 confirmed previous findings that area 1 is densely interconnected with VP. In addition, there appear to be sparse connections with VPS. There was no evidence of connections with Pa. Evidence from injection sites that extended from area 2 into areas 5 and 7, and from injection sites in area 5, indicates that the lateral posterior nucleus (LP) projects to rostral areas 5 and 7. The results support the conclusion that area 2 is a functionally distinct subdivision of somatosensory cortex, and indicate that area 2 has thalamic connections that are characteristic of both “sensory” (VP and VPS) and “association” (Pa) cortical fields.     

Topography of visual and somatosensory inputs to the pontine nuclei in zebra finches (Taeniopygia guttata)

Birds have a comprehensive network of sensorimotor projections extending from the forebrain and midbrain to the cerebellum via the pontine nuclei, but the organization of these circuits in the pons is not thoroughly described. Inputs to the pontine nuclei include two retinorecipient areas, nucleus lentiformis mesencephali (LM) and nucleus of the basal optic root (nBOR), which are important structures for analyzing optic flow. Other crucial regions for visuomotor control include the retinorecipient ventral lateral geniculate nucleus (GLv), and optic tectum (TeO). These visual areas, together with the somatosensory area of the anterior (rostral) Wulst, which is homologous to the primary somatosensory cortex in mammals, project to the medial and lateral pontine nuclei (PM, PL). In this study, we used injections of fluorescent tracers to study the organization of these visual and somatosensory inputs to the pontine nuclei in zebra finches. We found a topographic organization of inputs to PM and PL. The PM has a lateral subdivision that predominantly receives projections from the ipsilateral anterior Wulst. The medial PM receives bands of inputs from the ipsilateral GLv and the nucleus laminaris precommisulis, located medial to LM. We also found that the lateral PL receives a strong ipsilateral projection from TeO, while the medial PL and region between the PM and PL receive less prominent projections from nBOR, bilaterally. We discuss these results in the context of the organization of pontine inputs to the cerebellum and possible functional implications of diverse somato-motor and visuomotor inputs and parcellation in the pontine nuclei.     

Connections of the posterior thalamic region with the auditory ectosylvian cortex in the dog

The purpose of the present study was to define auditory cortical areas in the dog on the basis of thalamocortical connectivity patterns. Connections between the posterior thalamic region and auditory ectosylvian cortex were studied using axonally transported tracers: fluorochromes and biotinylated dextran amine. Cyto- and chemoarchitecture provided grounds for the division of the posterior thalamic region into three complexes, medial geniculate body (MGB), posterior nuclei (Po), and lateromedial and suprageniculate nuclei (LM-Sg). Distinctive cytoarchitectonic features and the distribution of dominant thalamocortical connections (determined quantitatively) allowed us to define four ectosylvian areas: middle (EM), anterior (EA), posterior (EP), and composite (CE). We found that each area was a place of convergence for projections from five to eleven nuclei of the three thalamic complexes, with dominant projections derived from one or two nuclei. Dominant topographical projections from the ventral nucleus to area EM confirmed physiological reports that it may be considered a primary auditory area (AI). We found the anterior part of the EM to be distinct in having unique strong connections with the deep dorsal MGB nucleus. Area EA, which receives dominant projections from the lateral Po (Pol) and medial MGB nuclei, as well as area EP, which receives dominant connections from the dorsal caudal MGB nucleus, compose two parasensory areas. Area CE receives dominant projections from the extrageniculate nuclei, anterior region of the LM-Sg, and Pol, supplemented with an input from the somatosensory VP complex, and may be considered a polymodal association area.     

Thalamic and corticocortical connections of the second somatic sensory area of the mouse

Thalamic and corticocortical connections of the second somatic sensory area (SII) in the mouse cerebral cortex were investigated by means of the retrograde transport of horseradish peroxidase. Focal injections of the enzyme were made in physiologically determined locations within the parietal cortex. Results show that SII receives substantial inputs from topographically appropriate regions within the ipsilateral ventrobasal nucleus and from the ipsilateral posterior group. The limb representation, which was previously found to be responsive to auditory stimulation, received inputs also from the medial division of the medial geniculate body. The SII face representation, which is largely unresponsive to auditory stimuli, received little or no input from the medial geniculate body. SII injections yielded retrograde labeling in the topographically appropriate region in the first somatic sensory area (SI), and SI injections retrogradely labeled cells in SII in a pattern consistent with previous electrophysiological maps. Homotypical regions within SI and SII therefore appear to be reciprocally interconnected. SII also receives inputs from the ipsilateral motor cortex and from contralateral SI and SII. Finally, injections into the SI paw but not face regions yielded retrograde labeling in the thalamic ventrolateral nucleus. Thus, the distal limb representations in SI and SII each receive inputs from a third major relay nucleus (i.e., medial geniculate to SII, ventrolateral nucleus to SI) whereas the face representations do not. These results indicate a close functional interrelationship between homotypical areas in SI and SII, though the two areas differ in several important respects. It is proposed that SII in mice may complement the function of SI by helping to define the overall sensory context in which detailed tactile discriminations are made.     

Anatomical and functional organization of somatosensory areas of the lateral fissure of the New World titi monkey (Callicebus moloch)

The organization of anterior and lateral somatosensory cortex was investigated in titi monkeys (Callicebus moloch). Multiunit microelectrode recordings were used to identify multiple representations of the body, and anatomical tracer injections were used to reveal connections. (1) Representations of the face were identified in areas 3a, 3b, 1, S2, and the parietal ventral area (PV). In area 3b, the face was represented from chin/lower lip to upper lip and neck/upper face in a rostrocaudal sequence. The representation of the face in area 1 mirrored that of area 3b. Another face representation was located in area 3a. Adjoining face representations in S2 and PV exhibited mirror-image patterns to those of areas 3b and 1. (2) Two representations of the body, the rostral and caudal ventral somatosensory areas (VSr and VSc), were found in the dorsal part of the insula. VSc was roughly a reversal image of the S2 body representation, and VSr was roughly a reversal of PV. (3) Neurons in the insula next to VSr and VSc responded to auditory stimuli or to both auditory and somatosensory stimuli. (4) Injections of tracers within the hand representations in areas 3b, 1, and S2 revealed reciprocal connections between these three areas. Injections in areas 3b and 1 labeled the ventroposterior nucleus, whereas injections in S2 labeled the inferior ventroposterior nucleus. The present study demonstrates features of somatosensory cortex of other monkeys in titi monkeys, while revealing additional features that likely apply to other primates.     

Connections of the lateral and medial divisions of the goldfish telencephalic pallium

Biotinylated dextran amine and fluorescent carbocyanine dye (DiI) were used to examine connections of the lateral (Dl) and medial (Dm) divisions of the goldfish pallium. Besides numerous intrinsic telencephalic connections to Dl and Dm, major ascending projections to these pallial divisions arise in the preglomerular complex of the posterior tuberculum, rather than in the dorsal thalamus. The rostral subnucleus of the lateral preglomerular nucleus receives auditory input via the medial pretoral nucleus, lateral line input via the ventrolateral toral nucleus, and visual input via the optic tectum, and it projects to both Dl and Dm. The anterior preglomerular nucleus and caudal subnucleus of the lateral preglomerular nucleus receive auditory input via the central toral nucleus and project to Dm. This pallial division also receives chemosensory information via the medial preglomerular nucleus. The central posterior (CP) nucleus, which receives both auditory and visual inputs, also projects to Dm and is the only dorsal thalamic nucleus projecting to the pallium. Thus, both Dl and Dm clearly receive multisensory inputs. Major projections of CP and projections of all other dorsal thalamic nuclei are to the subpallium, however. Descending projections of Dl are primarily to the preoptic area and the caudal hypothalamus, whereas descending projections of Dm are more extensive and particularly heavy to the anterior tuber and nucleus diffusus of the hypothalamus. The topography and connections of Dl are remarkably similar to those of the hippocampus of tetrapods, whereas the topography and connections of Dm are similar to those of the amygdala.     

Connections of the parahippocampal cortex in the cat. II. Subcortical afferents

The organization of subcortical inputs to the parahippocampal cortex, which in the present study in the cat is considered to comprise the entorhinal and perirhinal cortices, was studied by using retrograde and anterograde tracing techniques. The results of the retrograde tracer horseradish peroxidase (HRP), HRP conjugated with wheat germ agglutinine (WGA-HRP), Fast Blue (FB) or Nuclear Yellow (NY] injections indicate that the entorhinal and perirhinal cortices receive inputs from the magnocellular basal forebrain and from distinct portions of the amygdaloid complex, the claustrum, and the thalamus. The two cortices are further projected upon by fibers from the supramamillary region of the hypothalamus, the ventral tegmental area of the mesencephalon, the dorsal raphe nucleus, the nucleus centralis superior, and the locus coeruleus. The entorhinal cortex, in addition, receives projections from the medial septum. As regards the projections from the amygdaloid complex, it was observed that the entorhinal cortex receives its heaviest input from the basolateral amygdaloid nucleus, whereas the perirhinal cortex receives a strong projection from the lateral nucleus and a weaker projection from the basomedial nucleus of the amygdala. Of the thalamic nuclei that project to the parahippocampal cortex, the nucleus reuniens is only connected with the entorhinal cortex, while fibers from the medial geniculate nucleus and the lateral posterior nucleus terminate in the perirhinal cortex. Injections of tritiated amino acid (3H-leucine) were placed in the medial septum, the dorsal and ventral claustrum, the basolateral and basomedial amygdaloid nuclei, and the nucleus reuniens of the thalamus. The results of these experiments demonstrate that, with the exception of the claustrum, these subcortical areas project mainly to the superficial layers I-III and the lamina dissecans of the parahippocampal cortex, and to a lesser degree to the deep layers V and VI.     

Comparing thalamocortical and corticothalamic microstructure and spatial reciprocity in the macaque ventral posterolateral nucleus (VPLc) and medial pulvinar.

The detailed morphology of thalamocortical (TC) and corticothalamic (CT) pathways connecting the ventral posterolateral nucleus (VPLc) with the primary somatosensory cortex (areas 3b and 1) and the thalamic pulvinar with the posterior parietal cortex (primarily area 7a), was compared. Each pathway processes information relevant to directed reaching tasks, but whereas VPLc receives its major input from the spinal cord and external environment, the primary afferent to the pulvinar is cortical. Using combined tracer and thick fixed slice procedures, the soma/dendritic morphology of TC neuron populations (with known destination) was shown to be quantitatively similar within VPLc and the pulvinar. This implies that differences in information processing in VPLc (a primary relay) and the pulvinar (an integrative thalamic nucleus) are not defined by a distinctive TC morphology, but rather by the connections of these neuron populations. Two morphologically distinct types of CT axon were observed within the medial pulvinar and VPLc. The more common "Type E" were fine, had boutons en passant and diffuse terminal bifurcations ending in masses of tiny boutons. "Type R" axons were thicker, smooth, and terminated in localised clusters of large terminal boutons. Each type had a unique pattern of termination reflecting a distinct action on target neuron populations. The spatial relationship between TC distribution territories and CT terminal fields was examined within the medial pulvinar and VPLc by using anterograde and retrograde tracers injected together within cortical areas 7a, and 3b/1, respectively. Spatial overlap was incomplete within both thalamic nuclei. Our findings show a more complex relationship between TC and CT neuron populations than previously demonstrated.     

Thalamo-cortical connections of areas 3a and M1 in marmoset monkeys.

The present investigation is part of a broader effort to examine cortical areas that contribute to manual dexterity, reaching, and grasping. In this study we examine the thalamic connections of electrophysiologically defined regions in area 3a and architectonically defined primary motor cortex (M1). Our studies demonstrate that area 3a receives input from nuclei associated with the somatosensory system: the superior, inferior, and lateral divisions of the ventral posterior complex (VPs, VPi, and VPl, respectively). Surprisingly, area 3a receives the majority of its input from thalamic nuclei associated with the motor system, posterior division of the ventral lateral nucleus of the thalamus (VL), the mediodorsal nucleus (MD), and intralaminar nuclei including the central lateral nucleus (CL) and the centre median nucleus (CM). In addition, sparse but consistent projections to area 3a are from the anterior pulvinar (Pla). Projections from the thalamus to the cortex immediately rostral to area 3a, in the architectonically defined M1, are predominantly from VL, VA, CL, and MD. There is a conspicuous absence of inputs from the nuclei associated with processing somatic inputs (VP complex). Our results indicate that area 3a is much like a motor area, in part because of its substantial connections with motor nuclei of the thalamus and motor areas of the neocortex (Huffman et al. [2000] Soc. Neurosci. Abstr. 25:1116). The indirect input from the cerebellum and basal ganglia via the ventral lateral nucleus of the thalamus supports its role in proprioception. Furthermore, the presence of input from somatosensory thalamic nuclei suggests that it plays an important role in somatosensory and motor integration.     

Distribution of neurons immunoreactive for parvalbumin and calbindin in the somatosensory thalamus of the raccoon

The aim of this study was to assess the distribution of neurons immunoreactive for parvalbumin (PV), calbindin (CaBP), glutamic acid decarboxylase (GAD), and γ-aminobutyric acid (GABA) in the somatosensory thalamus of the raccoon and to compare these features to those of other species, especially primates. Immunohistochemistry was used to study the location of these neurons in the ventroposterior nucleus (VP), ventroposterior inferior nucleus (VPI), posterior group of nuclei (Po), and reticular nucleus (Rt). A consistent differential pattern of PV-positive (PV⁺) and CaBP-positive (CaBP⁺) cells was found in the somatosensory thalamus. Many PV⁺ neurons were observed in VP and Rt, but very few were found in VPI or Po. In contrast, CaBP⁺ neurons were distributed throughout VP, VPI, and Po but were very sparse or absent in Rt. In the VP nucleus, PV⁺ cells were distributed in clusters separated by interclusteral regions with a sparse distribution of PV⁺ cell bodies. The distributions of PV⁺ and CaBP⁺ cells tended to be complementary to each other in VP; regions with a high density of PV⁺ neurons had a low density of CaBP⁺ cell bodies. Double-labeling experiments revealed very few neurons in which PV and CaBP immunoreactivities were colocalized. Cells immunoreactive for GAD or GABA were found in PV-dense clusters of VP; fewer GABAergic neurons were present in the CaBP-dense interclusteral regions of VP and in VPI and Po. GAD⁺ and GABA⁺ neurons were most prominently distributed in Rt. We conclude that the distributions of PV⁺ and CaBP⁺ cell bodies in the raccoon somatosensory thalamus are very similar to those in primates. The density of GABAergic neurons in the somatosensory thalamus of the raccoon is less than that in the cat and monkey, but the relative distribution of GABAergic neurons in the different somatosensory nuclei is very similar to that in the cat and monkey. These results are discussed in relation to findings in other species and are related to the functions of lemniscal and nonlemniscal somatosensory pathways. J. Comp. Neurol. 388:120–129, 1997. © 1997 Wiley-Liss, Inc.