Cytoarchitecture and thalamic afferents of the sylvian and composite posterior gyri of the canine temporal cortex

The composite posterior and sylvian gyri of the canine temporal cortex show cytoarchitectonic features of poorly differentiated isocortex. Quantitative evaluation of connections examined with retrogradely transported fluorescent tracers indicated that both gyri received strong thalamic projections from the medial geniculate body (MG) and the lateromedial-suprageniculate (LM-Sg) complex, and a weaker projection from the posterior (Po) nuclei. On the basis of the connectivity patterns and cytoarchitectonic features we distinguished the anterior (CPa) and posterior (CPp) areas in posterior composite gyrus and the anterior (SA), dorsal (SD) and posterior (SP) sylvian areas. Afferents from individual thalamic nuclei were focused in distinct areas, forming dominant projections, and diminished gradually in the adjacent areas as non-dominant projections. The most prominent MG projection arose from the dorsal caudal (MGdc) nucleus. Its ventral subdivision sent a dominant projection into the SP and CPa, whereas the dorsal MGdc subdivision was connected with the SA, SD and CPp areas. The most substantial connections from the LM-Sg complex were directed to areas SA, SD and CPp, with weak connections to areas CPa and SP. A gradient of density of LM-Sg afferents was distributed in the opposite direction to that sent from the MGdc. The origin of the CPa and SP afferents in the ventral MGdc, like connections reaching the posterior ectosylvian cortex, suggest that these areas are related to processing of auditory information. In contrast, areas SA and CPp, receive dominant projections from the polymodal LM-Sg, and therefore may constitute successive steps in a hierarchy of cortical areas.     

Organization of cortical and subcortical projections to the feline insular visual area, IVA.

Extracellular recordings with carbon fiber-filled microelectrodes were used to identify the visually responsive area within the insular cortex (referred to hereafter as the insular visual area, IVA) of anaesthetized cats. Broadly speaking, IVA comprises the cortex surrounding the anterior ectosylvian sulcus (AEs) along its ventral bank and the major portion of the anterior sylvian gyrus. Visually sensitive cells were recorded along the whole length of the AEs. In the same animals, the afferent connections of IVA were studied through the use of the retrograde tracers wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) and fluorescent Diamidino yellow (DY), in combination with standard electrophysiological stimulation and recording techniques. The results indicate that: (1) the IVA receives a wide variety of telencephalic inputs, not only from visual, sensorimotor, auditory, limbic and association cortical areas, and from the claustrum, amygdala and basal nucleus of Meynert, as well, but also from the diencephalic projections arising mainly from the lateralis medialis-suprage niculate nuclear complex (LM-Sg) and the ventral medial nucleus (VM). (2) The gyral part of IVA (gIVA) receives afferents mainly from the lateral part of the lateral suprasylvian visual area (LS) throughout almost its entire length, as well as from area 20, the posterior suprasylvian sulcal area (PS), the frontal eye fields, areas 6 and 36, and almost the whole length of the cortical area lying along the anterior ectosylvian sulcus (AEs). (3) By contrast with (2), the sulcal part of IVA (sIVA) which corresponds to the anterior part of the anterior ectosylvian visual area (AEV) of Norita et al. ('86), receives cortical projections mainly from the lateral and medial parts of the anterior half of LS, area 20, PS, the frontal eye fields, area 36, and most parts of the cortical area extending along the AEs. (4) Subcortically, IVA receives thalamic afferents mainly from VM and LM-Sg. The connections between IVA and LM-Sg are organized topographically, with the more anterior part of IVA being related to the more ventral portion of LM-Sg, and with sIVA being related chiefly to the mid-portions of LM-Sg. These results thus suggest that IVA may function as an integrative centre among structures belonging to the extrageniculostriate system, the sensorimotor system, as well as to the limbic system. Furthermore, our electrophysiological and anatomical findings, together with previous reports concerning AEV, suggest that the posterior part of AEV (AEV proper) is distinctive from gIVA, and that the sIVA apparently serves as a transitional region between AEV and gIVA.     

Thalamic projections to the second and fourth somesthetic areas in the anterior ectosylvian gyrus of the cat

The topical organization of thalamic projections to the second and fourth somesthetic areas in the anterior ectosylvian gyrus of the cat has been studied using the technique of retrograde axonal transport of horseradish peroxidase. The projections of the posterolateral and posteromedial ventral nuclei (VPL, VPM) to the second somesthetic area (SII) are organized somatotopically. The posterior portion of SII (hindlimb area) receives fibers mainly from the dorsolateral part of VPL, the middle portion of SII (forelimb area) from the ventromedial part of VPL, and the anterior portion of SII (face area) from VPM. These topical projections are more loosely organized and less densely arranged than those to the first somesthetic area. The SII receives a few fibers from the medial geniculate nucleus, particularly its magnocellular and dorsal principal parts, and from the suprageniculate nucleus. The posterior part of SII lying near the secondary auditory area receives many fibers from the medial geniculate and suprageniculate nuclei, and only a few fibers from the lateral central and paracentral nuclei. The fourth somesthetic area (SIV), located in the dorsal bank of the anterior ectosylvian sulcus, receives fibers mainly from the dorsal principal and magnocellular parts of the medial geniculate nucleus, and from the suprageniculate nucleus. The SIV receives a fair number of fibers from VPL and VPM roughly in a somatotopical manner. The posterior portion of SIV receives fibers chiefly from the dorsolateral part of VPL, the middle portion of SIV from the ventromedial part of VPL, and the anterior portion from VPM. In addition, SIV receives a few fibers from the lateral central, paracentral, ventral lateral and ventral medial nuclei. The SIV, together with the most posterior part of SII, forms an auditory area, receiving many fibers from the medial geniculate and suprageniculate nuclei, and a few fibers from the intralaminar nuclei.     

Visual and auditory association areas of the cat's posterior ectosylvian gyrus: thalamic afferents

The feline posterior ectosylvian gyrus contains a broad band of association cortex that is bounded anteriorly by tonotopic auditory areas and posteriorly by retinotopic visual areas. To characterize the possible functions of this cortex and to throw light on its pattern of internal divisions, we have carried out an analysis of its thalamic afferents. Deposits of differentiable retrograde tracers were placed at 17 cortical sites in nine cats. The deposit sites spanned the crown of the posterior ectosylvian gyrus and adjacent cortex in the suprasylvian sulcus. We compiled counts of retrogradely labeled neurons in 12 thalamic nuclei delineated by use of Nissl and acetylcholinesterase stains. We then employed a statistical clustering algorithm to identify groups of injections that gave rise to similar patterns of thalamic labeling. The results suggest that the posterior ectosylvian gyrus contains 3 fundamentally different cortical districts that have the form of parallel vertical bands. Very anterior cortex, overlapping previously identified tonotopic auditory areas (AI, P and VP) receives a dense projection from the laminated division of the medial geniculate body (MGl). An intermediate strip, to which we refer as the auditory belt, is innervated by axons from nontonotopic divisions of the medial geniculate body (MGds, MGvl, MGm, and MGd), from the lateral division of the posterior group (Pol), and from the posterior suprageniculate nucleus (SGp). A posterior strip, to which we refer as EPp, receives strong projections from the LM-SG complex (LM-SGa and LMp), and lighter projections from the intralaminar and lateroposterior (LPm and LPl) nuclei. On grounds of thalamic connectivity, EPp is not obviously distinguishable from adjacent retinotopic visual areas (PLLS, DLS, and VLS), and may be regarded as forming, together with these areas, a connectionally homogeneous visual belt.     

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 dorsal zone of cat auditory cortex

The present study examined the anatomic connections of the dorsal zone of cat auditory cortex (DZ). The DZ was discriminated physiologically from the primary auditory field (AI) on the basis of neuronal responses with long latency and broad or multipeaked tuning curves. Wheat germ agglutinin-horseradish peroxidase was then injected either by pressure or iontophoretically. The thalamocortical and corticothalamic connections of the DZ were visualized by the presence of retrogradely labeled neurons and anterogradely labeled terminal fields in the thalamus; ipsilateral corticocortical projections from other cortical fields were visualized by the presence of retrogradely labeled cells. Injections of tracer into the DZ retrogradely labeled cells mainly in the lateral division of posterior complex (Po) and in the dorsal division (MGd) of the medial geniculate body (MGB); fewer labeled cells were found in the ventral (MGv) and medial (MGm) divisions of the MGB and in the suprageniculate nucleus. The DZ projection to Po, MGv, and MGd was heavy and was more diffuse than the reciprocal thalamocortical projection; the projection to MGm was light. The corticothalamic terminations and thalamocortical cells projecting to the same part of the DZ were not superimposed rigidly. The DZ received cortical projections from AI and from the second, anterior, and posterior auditory fields, and there were strong intra-DZ connections. Together with the physiological findings, the present results suggest that the DZ is a potentially separate auditory field from AI and is likely to be involved in both temporal and spectral integration of acoustic information. J. Comp. Neurol. 400:334–348, 1998. © 1998 Wiley-Liss, Inc.     

Organization of connections underlying the processing of auditory information in the dog

1. The canine temporal cortex includes the ectosylvian, composite posterior and sylvian gyri. 2. The distinctive features of the canine temporal cortex include the ectosylvian sulcus closed in its dorsal side and the substantial development of neocortex located within the posterior composite gyrus. 3. Thalamofugal connections from particular nuclei of the medial geniculate body, posterior thalamus and lateromedial-suprageniculate complex project to specific areas of the canine temporal cortex and are arranged as dominant and non-dominant projections. 4. Local intracortical connections distinguish the ectosylvian and posterior composite areas as unimodal auditory cortex. Long distant connections and polymodal convergence indicate that the composite ectosylvian area of the anterior ectosylvian gyrus and the anterodorsal sylvian areas are higher order association cortex. 5. Analysis of both thalamo-cortical and intracortical connections indicate that auditory processing in the cortex occurs in successive, hierarchically organized stages and in two main, anterior and ventral pathways.     

Differential thalamic connections of the posteroventral and dorsal posterior cingulate gyrus in the monkey

Previous functional studies suggest that the posterior cingulate gyrus is involved in spatial memory and its posteroventral part, in particular, is also involved in auditory memory. However, it is not clear whether the neural connections of the posteroventral part differ from those of the rest of the posterior cingulate gyrus. Here, we describe the thalamic connections of the posteroventral part of monkey area 23b (pv-area 23b), the main component of the posteroventral posterior cingulate gyrus. We compare these thalamic connections with those of the more dorsal area 23b (d-area 23b) and of adjoining retrosplenial areas 29 and 30. Thalamocortical projections to pv-area 23b originate mainly from the anterior nuclei, nucleus lateralis posterior and medial pulvinar. In contrast, projections to d-area 23b originate from the nucleus lateralis posterior, medial pulvinar, nucleus centralis latocellularis, mediodorsal nucleus and nucleus ventralis anterior and lateralis and weakly from the anterior nuclei. Projections to retrosplenial areas 29 and 30 originate from the anterior nuclei. Corticothalamic projections from pv-area 23b terminate in the anterior and laterodorsal nuclei, nucleus lateralis posterior and medial pulvinar. Projections from d-area 23b terminate in these nuclei as well as the nucleus ventralis anterior and lateralis. Projections from area 30 terminate mainly in the anterior nuclei and, to a lesser extent, in the medial pulvinar. These results show that the connections of pv-area 23b differ from those of d-area 23b or areas 29 and 30. This suggests that pv-area 23b may play distinct functional roles in memory processes, such as spatial and auditory memory.     

Thalamocortical connections of anterior and posterior parietal cortical areas in New World titi monkeys

We examined the thalamocortical connections of electrophysiologically identified locations in the hand and forelimb representations in areas 3b, 1, and 5 in the New World titi monkeys (Callicebus moloch), and of area 7b/AIP. Labeled cells and terminals in the thalamus resulting from the injections were related to architectonic boundaries. As in previous studies in primates, the hand representation of area 3b has dense, restricted projections predominantly from the lateral division of the ventral posterior nucleus (VPl). Projections to area 1 were highly convergent from several thalamic nuclei including the ventral lateral nucleus (VL), anterior pulvinar (PA), VPl, and the superior division of the ventral posterior nucleus (VPs). In cortex immediately caudal to area 1, what we term area 5, thalamocortical connections were also highly convergent and predominantly from nuclei of the thalamus associated with motor, visual, or somatic processing such as VL, the medial pulvinar (PM), and PA, respectively; with moderate projections from VP, central lateral nucleus (CL), lateral posterior nucleus (LP), and VPs. Finally, thalamocortical connections of area 7b/AIP were from a range of nuclei including PA, PM, LP/LD, VL, CL, PL, and CM. The current data support two conclusions drawn from previous studies in titi monkeys and other primates. First, cortex caudal to area 1 in New World monkeys is more like area 5 than area 2. Second, the presence of thalamic input to area 5 from both motor nuclei and somatosensory nuclei of the thalamus, suggests that area 5 could be considered a highly specialized sensorimotor area.     

Afferent connections of the lateralis medialis thalamic nucleus in the cat

We have used anterograde and retrograde horseradish peroxidass tracing methods in this study. Peroxidase injections in the lateralis medialis thalamic nucleus (LIB of the cat resulted in labeled neurons in cortical and subcortical structures that averaged 71 % and 29%, respectively. Every LM sector receives abundant projections from the polymodal sylvian anterior cortical area, the reticular thalamic nucleus, and the stratum opticum and intermediate layer of the superior colliculus. Less abundant but consistent projections were detected in cingular, auditory II, lateral suprasylvian and anterior ectosylvian visual cortices, and cortical area 7. A topographical distribution of afferent connections to different LM sectors arising from other cortical and subcortical structures could be established. The ventromedial sector receives connections mainly from the insular agranular, limbic and prefrontal cortical areas, as well as from brain stem structures and the contralateral pretectal region. The dorsolateral sector is mainly related to cortical areas and subcortical strictures processing visual information. The existence of overlap among neuronal LM populations receiving and sending connections to and from various cortical areas suggests that this nucleus is an appropriate substrate for effective interaction between different and distant cortical areas.     

Connections between the thalamus and the somatosensory areas of the anterior ectosylvian gyrus in the cat

The thalamocortical and corticothalamic connections of the second somatic sensory area (SII) and adjacent cortical areas in the cat were studied with anterograde and retrograde tracers. Injections consisted of horseradish peroxidase conjugated to wheat germ agglutinin (HRP-WGA) or a mixture of equal parts of tritiated leucine and proline. The cortical regions to be injected were electrophysiologically studied with microelectrodes to determine the localization of the selected components of the body representation in SII. The distribution of recording points was correlated in each case with the extent of the injection mass in the cortex. Distributions of retrograde and anterograde labeling in the thalamus were reconstructed from serial coronal sections. The results from cases with injections of tracers exclusively confined to separate parts of the body map in SII indicated a fairly precise topographical organization of projections from the ventrobasal complex (VB) to SII. The labeled cells and fibers were located within a series of lamella-like rods that curved throughout the dorsoventral and rostrocaudal axis of VB. The position and extent of these lamellae shifted from medial and ventral, in the medial subdivision of ventral posterior lateral nucleus (VPLm) for radial forelimb digit zones of SII, to dorsal, Posterior, and lateral, in the lateral subdivision of ventral posterior lateral nucleus (VPLl) for proximal leg and trunk regions in SII. For every injected area in SII the densest clustering of labeled cells and fibers was usually more posteriorly represented in VB. The distribution in these dense zones of labeling often extended through the central core of VB. SII projecting neurons were also consistently noted in the extreme rostral portion of the medial subdivision of the posterior nuclei (Pom) that lies dorsal to VB. Corticothalamic and thalamocortical connections for SII Were entirely reciprocal. Injections of tracers into cortical areas surrounding SII labeled other parts of the posterior complex but failed to label any part of VB except when the injection mass also diffused into SII. Injections into the somatic sensory cortex located lateral to SII, within the lips and depth of the upper bank of the anterior ectosylvian sulcus (AES), heavily labeled the central and posterior portions of Pom. Substantial labeling was noted in the lateral (Pol) and intermediate (Poi) divisions of Po only when the injections involved some part of the auditory area that occupies the most posterior part of the AEG and both banks of the immediately adjoining AES. The magnocellular nucleus of the medial geniculate (MGmc) was labeled only when some part of the auditory cortex was injected. The suprageniculate nucleus (SG) was labeled from the insula and lower bank of the AES. These results indicated that medial (rostral and caudal Pom) and lateral components (Poi, Pol, MGmc) of the Posterior complex have separate cortical projection zones to somatic sensory and auditory cortical regions, respectively. SIV and the lateral extent of area 5a located in the medial bank of the anterior suprasylvian sulcus sent projections to the deep layers of the supe- rior colliculus and the ventrolateral periaqueductal gray. No cortico-tectal projections were seen from SII.     

Telencephalic connections in lizards. II. Projections to anterior dorsal ventricular ridge

Three distinct cytoarchitectonic regions were identified within the anterior dorsal ventricular ridge (ADVR) of two species of lizards, Gekko gecko and Iguana iguana. These regions have been named according to their general topographical positions: medial area, caudolateral area, and rostrolateral area. Injections of horseradish peroxidase throughout the ADVR demonstrated that each of the three areas of the ADVR receives projections from specific thalamic nuclei which are associated with specific sensory modalities. The medial area receives an auditory thalamic projection from nucleus medialis. The caudolateral area receives thalamic projections from nucleus medialis posterior and nucleus posterocentralis. The latter two nuclei were shown to receive projections from the spinal cord and, therefore, are presumed to be associated with body somatosensory information. The rostrolateral area receives a thalamic projection from nucleus rotundus, which receives visual information. In addition, the mesencephalic tegmentum and the thalamic nucleus dorsomedialis project to the entire ADVR. The latter projection is similar to the diffuse cortical projections of the intralaminar thalamic nuclei in mammals. These findings support previous suggestions that the ADVR is comparable to sensory regions of the mammalian neocortex.     

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.     

Projections of auditory cortex to the medial geniculate body of the cat

The corticofugal projection from 12 auditory cortical fields onto the medial geniculate body was investigated in adult cats by using wheat germ agglutinin conjugated to horseradish peroxidase or biotinylated dextran amines. The chief goals were to determine the degree of divergence from single cortical fields, the pattern of convergence from several fields onto a single nucleus, the extent of reciprocal relations between corticothalamic and thalamocortical connections, and to contrast and compare the patterns of auditory corticogeniculate projections with corticofugal input to the inferior colliculus. The main findings were that (1) single areas showed a wide range of divergence, projecting to as few as 5, and to as many as 15, thalamic nuclei; (2) most nuclei received projections from approximately five cortical areas, whereas others were the target of as few as three areas; (3) there was global corticothalamic-thalamocortical reciprocity in every experiment, and there were also significant instances of nonreciprocal projections, with the corticothalamic input often more extensive; (4) the corticothalamic projection was far stronger and more divergent than the corticocollicular projection from the same areas, suggesting that the thalamus and the inferior colliculus receive differential degrees of corticofugal control; (5) cochleotopically organized areas had fewer corticothalamic projections than fields in which tonotopy was not a primary feature; and (6) all corticothalamic projections were topographic, focal, and clustered, indicating that areas with limited cochleotopic organization still have some internal spatial arrangement. The areas with the most divergent corticothalamic projections were polysensory regions in the posterior ectosylvian gyrus. The projection patterns were indistinguishable for the two tracers. These findings suggest that every auditory thalamic nucleus is under some degree of descending control. Many of the projections preserve the relations between cochleotopically organized thalamic and auditory areas, and suggest topographic relations between nontonotopic areas and nuclei. The collective size of the corticothalamic system suggests that both lemniscal and extralemniscal auditory thalamic nuclei receive significant corticofugal input.     

Thalamic connections of the auditory cortex in marmoset monkeys: core and medial belt regions

In this study and its companion, the cortical and subcortical connections of the medial belt region of the marmoset monkey auditory cortex were compared with the core region. The main objective was to document anatomical features that account for functional differences observed between areas. Injections of retrograde and bi-directional anatomical tracers targeted two core areas (A1 and R), and two medial belt areas (rostromedial [RM] and caudomedial [CM]). Topographically distinct patterns of connections were revealed among subdivisions of the medial geniculate complex (MGC) and multisensory thalamic nuclei, including the suprageniculate (Sg), limitans (Lim), medial pulvinar (PM), and posterior nucleus (Po). The dominant thalamic projection to the CM was the anterior dorsal division (MGad) of the MGC, whereas the posterior dorsal division (MGpd) targeted RM. CM also had substantial input from multisensory nuclei, especially the magnocellular division (MGm) of the MGC. RM had weak multisensory connections. Corticotectal projections of both RM and CM targeted the dorsomedial quadrant of the inferior colliculus, whereas the CM projection also included a pericentral extension around the ventromedial and lateral portion of the central nucleus. Areas A1 and R were characterized by focal topographic connections within the ventral division (MGv) of the MGC, reflecting the tonotopic organization of both core areas. The results indicate that parallel subcortical pathways target the core and medial belt regions and that RM and CM represent functionally distinct areas within the medial belt auditory cortex.     

The organization of thalamic connections

Connections ascending to the thalamus. Contrary to classical opinion, all thalamic nuclei receive extrathalamic afferents. Segregation or convergence within a topographically defined nucleus represent two modalities of thalamic afferents. In addition, certain topographically organized thalamic afferents possess "privileged" or primary "targets" in the thalamic nucleus while others possess supplementary "targets" in other thalamic nuclei (see cerebellar, pallidal and spinothalamic projections). Ascending connections from several brain stem structures can converge on the same nucleus or diverge to several thalamic nuclei. Thalamic connections with the telencephalon. Methods for determining axonal transport have demonstrated that all thalamic nuclei, with the exception of the reticular nucleus and the ventral part of the lateral geniculate body, project towards the cerebral cortex. Four nuclear complexes can be recognized in the cat as a function of the different modalities of localization, concentration and lamination of the projections towards the cortex and the central grey nuclei. In general, the thalamocortical connections have reciprocal ipsilateral corticothalamic projections originating in the infragranular layers of the cerebral cortex. The reticular nucleus and the ventral part of the lateral geniculate body, which is not projected to the cerebral cortex, are exceptions. Each cortical area receives a "privileged" connection from a thalamic nucleus and a supplementary connection- from one or several other thalamic nuclei. The "privileged" connections usually pass to the fourth and third layers of the neocortex, and sometimes also to the first layer. In contrast, the supplementary connections pass to different superficial or deep cortical layers. Each nucleus is formed of subunits which possess different hodologic and topographic characteristics as a function of the nucleus considered. Convergence or divergence of thalamocortical and corticothalamic projections on the different thalamic nuclei, as well as the laminar distribution of efferents in the cerebral cortex, are related strictly to the hodologic organization of different cellular subunits constituting the nuclei. Concentration or diffusion of thalamic projections on cerebral cortex is related more to the single or multiple projection of cell populations belonging to a thalamic nucleus than to widespread collateralization of thalamocortical axons.     

Connections of cat auditory cortex: I. Thalamocortical system

Despite the functional importance of the medial geniculate body (MGB) in normal hearing, many aspects of its projections to auditory cortex are unknown. We analyzed the MGB projections to 13 auditory areas in the cat using two retrograde tracers to investigate thalamocortical nuclear origins, topography, convergence, and divergence. MGB divisions and auditory cortex areas were defined independently of the connectional results using architectonic, histochemical, and immunocytochemical criteria. Each auditory cortex area received a unique pattern of input from several MGB nuclei, and these patterns of input identify four groups of cortical areas distinguished by their putative functional affiliations: tonotopic, nontonotopic, multisensory, and limbic. Each family of areas received projections from a functionally related set of MGB nuclei; some nuclei project to only a few areas (e.g., the MGB ventral division to tonotopic areas), and others project to all areas (e.g., the medial division input to every auditory cortical area and to other regions). Projections to tonotopic areas had fewer nuclear origins than those to multisensory or limbic-affiliated fields. All projections were organized topographically, even those from nontonotopic nuclei. The few divergent neurons (mean: 2%) are consistent with a model of multiple segregated streams ascending to auditory cortex. The expanded cortical representation of MGB auditory, multisensory, and limbic affiliated streams appears to be a primary facet of forebrain auditory function. The emergence of several auditory cortex representations of characteristic frequency may be a functional multiplication of the more limited maps in the MGB. This expansion suggests emergent cortical roles consistent with the divergence of thalamocortical connections.     

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)     

Thalamic connectivity of the second somatosensory area and neighboring somatosensory fields of the lateral sulcus of the macaque

The thalamocortical relations of the somatic fields in and around the lateral sulcus of the macaque were studied following cortical injections of tritated amino acids and horseradish peroxidase (HRP). Special attention was paid to the second somatosensory area (S2), the connections of which were also studied by means of thalamic isotope injections and retrograde degeneration. S2 was shown to receive its major thalamic input from the ventroposterior inferior thalamic nucleus (VPI) and not, as previously reported, from the caudal division of the ventroposterior lateral nucleus (VPLc). Following small injections of isotope or HRP into the hand representation of S2, only VPI was labeled. Larger injections, which included the representations of more body parts, led to heavy label in VPI, with scattered label in VPLc, the central lateral nucleus (CL), and the posterior nucleus (Po). In addition, small isotope injections into VPLc did not result in label in S2 unless VPI was also involved in the injection site, and ablations of S2 led to cell loss in VPI. Comparison of injections involving different body parts in S2 suggested a somatotopic arrangement within VPI such that the trunk and lower limb representations are located posterolaterally and the hand and arm representations anteromedially. The location of the thalamic representations of the head, face, and intraoral structures that project to S2 may be in the ventroposterior medial nucleus (VPM). The granular (Ig) and dysgranular (Id) fields of the insula and the retroinsular field (Ri) each receive inputs from a variety of nuclei located at the posteroventral border of the thalamus. Ig receives its heaviest input from the suprageniculate-limitans complex (SG-Li), with additional inputs from Po, the magnocellular division of the medial geniculate n. (MGmc), VPI, and the medial pulvinar (Pulm). Id receives its heaviest input from the basal ventromedial n. (VMb), with additional inputs from VPI, Po, SG-Li, MGmc, and Pulm. Ri receives its heaviest input from Po, with additional input from SG-Li, MGmc, Pulm, and perhaps VPI. Area 7b receives its input from Pulm, the oral division of the pulvinar, the lateral posterior n., the medial dorsal n., and the caudal division of the ventrolateral n. These results indicate that the somatic cortical fields, except for those comprising the first somatosensory area, each receive inputs from an array of thalamic nuclei, rather than just one, and that individual thalamic somatosensory relay nuclei each project to more than one cortical field.     

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.