An analysis of potentially converging inputs to the rostral ventral thalamic nuclei of the cat

Summary Potentially convergent inputs to cerebellar-receiving and basal ganglia-receiving areas of the thalamus were identified using horseradish peroxidase (HRP) retrograde tracing techniques. HRP was deposited iontophoretically into the ventroanterior (VA), ventromedial (VM), and ventrolateral (VL) thalamic nuclei in the cat. The relative numbers of labeled neurons in the basal ganglia and the cerebellar nuclei were used to assess the extent to which the injection was in cerebellar-receiving or basal ganglia-receiving portions of thalamus. The rostral pole of VA showed reciprocal connections with prefrontal portions of the cerebral cortex. Only the basal ganglia and the hypothalamus provided non-thalamic input to modulate these cortico-thalamo-cortical loops. In VM, there were reciprocal connections with prefrontal, premotor, and insular areas of the cerebral cortex. The basal ganglia (especially the substantia nigra), and to a lesser extent, the posterior and ventral portions of the deep cerebellar nuclei, provided input to VM and may modulate these corticothalamo-cortical loops. The premotor cortical areas connected to VM include those associated with eye movements, and afferents from the superior colliculus, a region of documented importance in oculomotor control, also were labeled by injections into VM. The dorsolateral portion of the VA-VL complex primarily showed reciprocal connections with the medial premotor (area 6) cortex. Basal ganglia and cerebellar afferents both may modulate this cortico-thalamo-cortical loop, although they do not necessarily converge on the same thalamic neurons. The cerebellar input to dorsolateral VA-VL was from posterior and ventral portions of the cerebellar nuclei, and the major potential brainstem afferents to this region of thalamus were from the pretectum. Mid- and caudo-lateral portions of VL had reciprocal connections with primary motor cortex (area 4). The dorsal and anterior portions of the cerebellar nuclei had a dominant input to this corticothalamo-cortical loop. Potentially converging brainstem afferents to this portion of VL were from the pretectum, especially pretectal areas to which somatosensory afferents project.List of Abbreviations AC central amygdaloid nucleus - AL lateral amygdaloid nucleus - AM anteromedial thalamic nucleus - AV anteroventral thalamic nucleus - BC brachium conjunctivum - BIC brachium of the inferior colliculus - Cd caudate nucleus - CL centrolateral thalamic nucleus - CM centre median nucleus - CP cerebral peduncle - CUN cuneate nucleus - DBC decussation of the brachium conjunctivum - DR dorsal raphe nuclei - EC external cuneate nucleus - ENTO entopeduncular nucleus - FN fastigial nucleus - FX fornix - GP globus pallidus - GR gracile nucleus - IC internal capsule - ICP inferior cerebellar peduncle - IP interpeduncular nucleus - IVN inferior vestibular nucleus - LD lateral dorsal thalamic nucleus - LGN lateral geniculate nucleus - LH lateral hypothalamus - LP lateral posterior thalamic complex - LRN lateral reticular nucleus - LVN lateral vestibular nucleus - MB mammillary body - MD mediodorsal thalamic nucleus - MG medial geniculate nucleus - ML medial lemniscus - MLF medial lengitudinal fasciculus - MT mammillothalamic tract - MVN medial vestibular nucleus - NDBB nucleus of the diagonal band of Broca - NIA anterior nucleus interpositus - NIP posterior nucleus interpositus - OD optic decussation - OT optic tract - PAC paracentral thalamic nucleus - PPN pedunculopontine region - PRO gyrus proreus - PRT pretectal region - PT pyramidal tract - PTA anterior pretectal region - PTM medial pretectal region - PTO olivary pretectal nucleus - PTP poterior pretectal region - Pul pulvinar nucleus - Put putamen - RF reticular formation - RN red nucleus - Rt reticular complex of the thalamus - S solitary tract - SCi superior colliculus, intermediate gray - SN substantia nigra - ST subthalamic nucleus - VA ventroanterior thalamic nucleus - VB ventrobasal complex - VL ventrolateral thalamic nucleus - VM ventromedial thalamic nucleus - III oculomotor nucleus - IIIn oculomotor nerve - 5S spinal trigeminal nucleus - 5T spinal trigeminal tract - VII facial nucleus     

An autoradiographic study of topographical relationships between pallidal and cerebellar projections to the cat thalamus

Summary Injections of ³H-leucine were made in the entopeduncular nucleus or dentate nucleus of the cerebellum in eight cats. The terminal projection zones of both pathways in the thalamus were studied using the sagittal plane and their relationships to one another as well as to cytoarchitectural boundaries of thalamic nuclei were compared. The data indicate that the territories controlled by the two projection systems are almost entirely segregated. The segregation is mainly along the antero-posterior axis as the main pallidal projection zone occupies the medio-ventral VA while the main dentate projection zone lies posterior to it in the VL. Furthermore, the dorsolateral part of the VA not occupied by pallidal projections receives dentate projections. In the VM, both afferent systems terminate in the lateral part of the nucleus with pallidal territory located anteriorly and dentate territory located posteriorly, again without overlap. As the delineations of nuclear subdivisions in the ventral thalamus of the cat have been a subject of some controversy, it is suggested that the boundaries of the VA, VL and VM in the cat thalamus be defined on the basis of basal ganglia and cerebellar projection zones.Abbreviations used in the Text and in Fig. 5 AM anterior medial nucleus - AV anterior ventral nucleus - BC brachium conjunctivum - CA anterior commissure - CC crus cerebri - CP posterior commissure - CD caudate nucleus - CE centrum medianum - CLN central lateral nucleus - DN dentate nucleus - EPN entopeduncular nucleus - FF Forel's field - FN fastigial nucleus - FR fasciculus retroflexus - HL lateral habenular nucleus - HM medial habenular nucleus - INA anterior interposite nucleus - INP posterior interposite nucleus - IC internal capsule - LD lateral dorsal nucleus - LG lateral geniculate body - MD medial dorsal nucleus - MTT mamillothalamic tract - NR red nucleus - OT optic tract - PAC paracentral nucleus - PF parafascicular nucleus - PV pulvinar - RT reticular thalamic nucleus - SM submedian nucleus - SN substantia nigra - SNr substantia nigra pars reticularis - STN subthalamic nucleus - VF ventral posterior nucleus - VA ventral anterior nucleus - VL ventral lateral nucleus - VM ventral medial nucleus - ZI zona incerta Supported in part by a grant from the American Parkinson Disease Association and NIH grant R01NS19280     

Cortical projections of the anterior thalamic nuclei in the cat

Summary Unilateral stereotaxic lesions were made in the anterior thalamic nuclei of the cat, and the ensuing terminal degeneration traced to the medial cortex by the methods of Nauta-Gygax and Fink-Heimer. The anterodorsal nucleus projects to the retrosplenial, postsubicular and presubicular areas. These projections appear to be organized in the dorsoventral direction. The posterior portion of the retrosplenial area receives no fibers from the anterodorsal nucleus. Fibers from this nucleus are distributed largely in layer I and in layer III and the deep portion of layer II of the posterior limbic cortex. The anteroventral nucleus sends fibers to the cingular area and parts of the retrosplenial, postsubicular and presubicular areas. These projections appear to be organized in a topical manner mediolaterally. When the lesion involves the parvocellular part of the nucleus, degeneration spreads to the lower lip, bank and fundus of the splenial sulcus. There appears to be an anteroposterior organization in the cortical projections of the anteroventral nucleus. Fibers from the anteroventral nucleus are distributed most profusely in layers IV and III and in the superficial portion of layer I of the posterior limbic cortex. The anteromedial nucleus sends fine fibers to the anterior limbic region and to the cingular, retrosplenial, postsubicular and presubicular areas. The cortical projections of the anteromedial nucleus appear to be topographically organized in the dorsoventral direction. Fibers from the anteromedial nucleus are distributed largely in cortical layers V and VI of the anterior and posterior limbic regions.Abbreviations used in Figures a anterior - AD anterodorsal nucleus - AM anteromedial nucleus - AMD dorsolateral part of anteromedial nucleus - AMV ventromedial part of anteromedial nucleus - AV anteroventral nucleus - AVM magnocellular part of anteroventral nucleus - AVP parvocellular part of anteroventral nucleus - CC corpus callosum - Cg cingular area - CM medial central nucleus - Il infralimbic area - LA anterior limbic region - LD dorsal lateral nucleus - MD dorsal medial nucleus - Of orbitofrontal region - p posterior - Pr presubicular area - Prag precentral agranular area - Ps postsubicular area - Pt paratenial nucleus - Pv anterior paraventricular nucleus - R reuniens nucleus - Rs retrosplenial area - Rt thalamic reticular nucleus - SC cruciate sulcus - SM stria medullaris - Sm submedial nucleus - SS splenial sulcus - VA ventral anterior nucleus - VL ventral lateral nucleus - VM ventral medial nucleus     

Connections of the anterior ectosylvian visual area (AEV)

Summary We have previously described a visual area situated in the cortex surrounding the deep infolding of the anterior ectosylvian sulcus of the cat (Mucke et al. 1982). Using orthograde and retrograde transport methods we now report anatomical evidence that this anterior ectosylvian visual area (AEV) is connected with a substantial number of both cortical and subcortical regions. The connections between AEV and other cortical areas are reciprocal and, at least in part, topographically organized: the rostral AEV is connected with the bottom region of the presylvian sulcus, the lower bank of the cruciate sulcus, the rostral part of the ventral bank of the splenial sulcus, the rostral portion of the lateral suprasylvian visual area (LS) and the lateral bank of the posterior rhinal sulcus; the caudal AEV is connected with the bottom region of the presylvian sulcus, the caudal part of LS, the ventral part of area 20 and the lateral bank of the posterior rhinal sulcus. Subcortically, AEV has reciprocal connections with the ventral medial thalamic nucleus (VM), with the medial part of the lateralis posterior nucleus (LPm), as well as with the lateralis medialis-suprageniculate nuclear (LM-Sg) complex. These connections are also topographically organized with more rostral parts of AEV being related to more ventral portions of the LPm and LM-Sg complex. AEV also projects to the caudate nucleus, the putamen, the lateral amygdaloid nucleus, the superior colliculus, and the pontine nuclei. It is concluded that AEV is a visual association area which functionally relates the visual with both the motor and the limbic system and that it might play a role in the animal's orienting and alerting behavior.Abbreviations Ac aqueductus cerebri - AEs anterior ectosylvian sulcus - ALLS anterolateral lateral suprasylvian area - AMLS anteromedial lateral suprasylvian area - ASs anterior suprasylvian sulcus - Cd caudate nucleus - CL central lateral nucleus - Cl claustrum - Cos coronal sulcus - Crs cruciate sulcus - DLS dorsal lateral suprasylvian area - GI stratum griseum intermediale - GP stratum griseum profundum - IC inferior colliculus - LAm lateral amygdaloid nucleus - LGNd dorsal nucleus of lateral geniculate body - LGNv ventral nucleus of lateral geniculate body - Llc nucleus lateralis intermedius, pars caudalis - LM nucleus lateralis medialis - LPl nucleus lateralis posterior, pars lateralis - LPm nucleus lateralis posterior, pars medialis - Ls lateral sulcus - MD nucleus mediodorsalis - MG medial geniculate body - MSs middle suprasylvian sulcus - Ndl nucleus dorsolateralis pontis - Nl nucleus lateralis pontis - Np nucleus peduncularis pontis - Npm nucleus paramedianus pontis - Nrt nucleus reticularis tegmenti pontis - Nv nucleus ventralis pontis - Ped cerebral peduncle - PEs posterior ectosylvian sulcus - Pg periaqueductal gray - PLLS posterolateral lateral suprasylvian area - PMLS posteromedial lateral suprasylvian area - PSs presylvian sulcus - Pul pulvinar - Put putamen - R red nucleus - Sg suprageniculate nucleus - SN substantia nigra - Sps splenial sulcus - Syls sylvian sulcus - T trapezoid body - VA ventral anterior nucleus - VL ventral lateral nucleus - VLS ventral lateral suprasylvian area - VM ventral medial nucleus - VPL ventral posterolateral nucleus - VPM ventral posteromedial nucleus Sponsored by Max-Planck-Society during part of the studySponsored by Thyssen FoundationSponsored by Alexander von Humboldt-Foundation     

Laminar and modular organization of prefrontal projections to multiple thalamic nuclei

The prefrontal cortex projects to many thalamic nuclei, in pathways associated with cognition, emotion, and action. We investigated how multiple projection systems to the thalamus are organized in prefrontal cortex after injection of distinct retrograde tracers in the principal mediodorsal (MD), the limbic anterior medial (AM), and the motor-related ventral anterior/ventral lateral (VA/VL) thalamic nuclei in rhesus monkeys. Neurons projecting to these nuclei were organized in interdigitated modules extending vertically within layers VI and V. Projection neurons were also organized in layers. The majority of projection neurons to MD or AM originated in layer VI (∼80%), but a significant proportion (∼20%) originated in layer V. In contrast, prefrontal neurons projecting to VA/VL were equally distributed in layers V and VI. Neurons directed to VA/VL occupied mostly the upper part of layer V, while neurons directed to MD or AM occupied mostly the deep part of layer V. The highest proportions of projection neurons in layer V to each nucleus were found in dorsal and medial prefrontal areas. The laminar organization of prefrontal cortico-thalamic projections differs from sensory systems, where projections originate predominantly or entirely from layer VI. Previous studies indicate that layer V cortico-thalamic neurons innervate through some large terminals thalamic neurons that project widely to superficial cortical layers. The large population of prefrontal projection neurons in layer V may drive thalamic neurons, triggering synchronization by recruiting several cortical areas through widespread thalamo-cortical projections to layer I. These pathways may underlie the synthesis of cognition, emotion and action.     

Segregation of lemniscal inputs and motor cortex outputs in cat ventral thalamic nuclei: application of a novel technique

Summary A double labeling method that permits accurate delineation of the terminals of medial lemniscal fibers was used to determine whether thalamic neurons projecting to motor cortex in the cat are in a position to be contacted by such terminals. Thalamic neurons in the VL nucleus were retrogradely labeled by injections of fluorogold placed in the cytoarchitectonically defined area 4, while lemniscal axons and their terminal boutons were anterogradely labeled, in a Golgi-like manner, from injections of Fast Blue placed under physiological control in different parts of the contralateral dorsal column nuclei. In additional experiments, spinothalamic fibers were similarly labeled by injections of Fast Blue in the spinal cord. The results reveal that there is no significant overlap in the distributions of lemniscal terminals and motor cortex-projecting neurons and that no somata or proximal dendrites of motor cortex-projecting neurons are in a position to receive lemniscal terminals. Spinothalamic terminals, on the other hand, end in clusters around motor cortex-projecting neurons in the VL nucleus as well as in other nuclei and are a more likely route for short latency somatosensory inputs to the motor cortex.Abbreviations AD anterodorsal nucleus - AM anteromedial nucleus - AP area postrema - AV anteroventral nucleus - C cuneate nucleus - CeM central medial nucleus - CL central lateral nucleus - CM centre médian nucleus - EC external cuneate nucleus - G gracile nucleus - L limitans nucleus - LD lateral dorsal nucleus - LP lateral posterior nucleus - MGM magnocellular medial geniculate nucleus - MD mediodorsal nucleus - MTT mamillothalamic tract - MV medioventral nucleus - Pc paracentral nucleus - Pf parafascicular nucleus - Po posterior nuclei - R reticular nucleus - RF fasciculus retroflexus - S solitary nucleus - SG suprageniculate nucleus - T spinal trigeminal nucleus - VA ventral anterior nucleus - VIN vestibular nuclei - VL ventral lateral nucleus - VMb basal ventral medial nucleus - VMp principal ventral medial nucleus - VPL ventral posterior lateral nucleus - VPM ventral posterior medial nucleus - ZI zona incerta - 1,2,3a,3b,4 fields of cerebral cortex - C4, C5, C6 spinal cord segments - 5SP,5ST spinal trigeminal nucleus and tract - 10, 12 vagal and hypoglossal nuclei     

The corticothalamic projection from the gyrus proreus and the medial wall of the rostral hemisphere in the cat an experimental study with silver impregnation methods

Summary The corticothalamic projections from the gyrus proreus and the medial wall of the rostral hemisphere have been studied in the cat with the silver method of Nauta. The gyrus proreus projects upon the following nuclei (for abbreviations, see list on page 133), ipsilateral R, VA, VM, VL, MD, Pc, CL, CM, Pf, VPM, VPMpc. VPI and to the contralateral principal nucleus of the trigeminal nerve. The medial wall of the rostral hemisphere projects bilaterally upon R, VA, VM, VL, MD, Pc, CL, CM, Pf, VPM, VPMpc, VPI, VPL, the dorsal column nuclei and the principal nucleus of the trigeminal nerve. The ipsilateral thalamic projection is more abundant than the contralateral. The latter appears to increase in amount as the lesion is placed successively more ventrally on the medial wall of the rostral hemisphere. Some degenerating fibers cross in the corpus callosum and descend in the contralateral internal capsule but the majority cross in the dorsal part of the anterior commissure and reach the medial aspect of the anterior limb of the contralateral internal capsule. A somatotopical organization of the medial wall of the rostral hemisphere has been demonstrated. The rostrocaudal part projects upon the ipsilateral VPL lateralis (VPLl) and nucleus cuneatus and the contralateral nucleus gracilis and VPL medialis (VPLm). The caudal part of this cortical area sends fibers bilaterally to VPM, VPMpc, and the principal nucleus of the trigeminal nerve. The intermediate part, which also includes agranular cortex on the medial wall, projects upon ispsilateral VPLm and nucleus gracilis and upon contralateral VPLl and nucleus cuneatus. — The fibers to the ventro-basal complex, dorsal column nuclei and the principal nucleus of the trigeminal nerve are rather thick. The corticofugal fibers to the other thalamic nuclei are quite thin. — The findings are discussed in light of relevant anatomical and physiological observations in the literature and special emphasis has been laid on reported observations on the supplementary motor area.     

An autoradiographic study of efferent connections of the globus pallidus in Macaca mulatta

Summary Radioactive amino acids were injected into restricted regions of the globus pallidus of rhesus macaques to allow identification of the organization and courses of efferent pallidal projections. The previously identified projection of the internal pallidal segment (GPi) to ventral thalamic nuclei showed a topographic organization, with the predominant projection from ventral GPi being to medial and caudal ventralis anterior (VA) and lateralis (VL) and from dorsal GPi to lateral and rostral VA and VL. Pallidal efferent fibers also extended caudally and dorsally into pars caudalis of VL, but they spared the portion of pars oralis of VL shown by others to receive input from the cerebellum. In addition to centromedian labeling in all animals, the parafascicular nucleus was also labeled when isotope was injected into dorsal GPi. The medial route from GPi to the midbrain tegmentum was more substantial than has been shown before, and along this route there was an indication that some fibers terminated in the prerubral region. The projection to the pedunculopontine nucleus was extensive, and fibers continued caudally into the parabrachial nuclei.Pallidal projections to the thalamus seem to be topographically organized but spare thalamic regions that interact with area 4. Caudally directed efferent fibers follow multiple routes and extend more caudally than to the pedunculopontine nuclei.Abbreviations Cd caudate nucleus - CM centromedian nucleus - CT central tegmental tract - DPCS decussation of superior cerebellar peduncle - F fornix - FLM medial longitudinal fasciculus - GPe globus pallidus, pars externa - GPi globus pallidus, pars interna - HbL lateral habenular nucleus - HbM medial habenular nucleus - Is interstitial nucleus - LM medial lemniscus - MD dorsomedial nucleus - PbL lateral parabrachial nucleus - PbM medial parabrachial nucleus - PCS superior cerebellar peduncle - Pf parafascicular nucleus - PPN pedunculopontine nucleus - Put putamen - R reticular nucleus - Rmg red nucleus, pars magnocellularis - Rpc red nucleus, pars parvocellularis - S stria medullaris - SI substantia innominata - SNc substantia nigra, pars compacta - SNr substantia nigra, pars reticulata - St subthalamic nucleus - ST stria terminalis - THI habenulointerpeduncular tract - TM tuberomamillary nucleus - TMT mamillothalamic tract - VA nucleus ventralis anterior - VAmg nucleus ventralis anterior, pars magnocellularis - VAp nucleus ventralis anterior, pars principalis - VI nucleus ventralis intermedius - VLc nucleus ventralis lateralis, pars caudalis - VLm nucleus ventralis lateralis, pars medialis - VLo nucleus ventralis lateralis, pars oralis - VPL nucleus ventralis posterior lateralis - X area X Supported by National Institutes of Health, grant RR00166, Rehabilitation Services Administration, grant 16-P-56818, and PHS grant NS10804     

Thalamic connections of the dorsal and ventral premotor areas in New World owl monkeys

Thalamic connections of two premotor cortex areas, dorsal (PMD) and ventral (PMV), were revealed in New World owl monkeys by injections of fluorescent dyes or wheat-germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). The injections were placed in the forelimb and eye-movement representations of PMD and in the forelimb representation of PMV as determined by microstimulation mapping. For comparison, injections were also placed in the forelimb representation of primary motor cortex (M1) of two owl monkeys. The results indicate that both PMD and PMV receive dense projections from the ventral lateral (VL) and ventral anterior (VA) thalamus, and sparser projections from the ventromedial (VM), mediodorsal (MD) and intralaminar (IL) nuclei. Labeled neurons in VL were concentrated in the anterior (VLa) and the medial (VLx) nuclei, with only a few labeled cells in the dorsal (VLd) and posterior (VLp) nuclei. In VA, labeled neurons were concentrated in the parvocellular division (VApc) dorsomedial to VLa. Labeled neurons in MD were concentrated in the most lateral and posterior parts of the nucleus. VApc projected more densely to PMD than PMV, especially to rostral PMD, whereas caudal PMD received stronger projections from neurons in VLx and VLa. VLd projected exclusively to PMD, and not to PMV. In addition, neurons labeled by PMD injections tended to be more dorsal in VL, IL, and MD than those labeled by PMV injections. The results indicate that both premotor areas receive indirect inputs from the cerebellum (via VLx, VLd and IL) and globus pallidus (via VLa, VApc, and MD). Comparisons of thalamic projections to premotor and M1 indicate that both regions receive strong projections from VLx and VLa, with the populations of cells projecting to M1 located more laterally in these nuclei. VApc, VLd, and MD project mainly to premotor areas, while VLp projects mainly to M1. Overall, the thalamic connectivity patterns of premotor cortex in New World owl monkeys are similar to those reported for Old World monkeys.     

Forebrain connections of medial agranular cortex in the prairie vole, Microtus ochrogaster

 Fluorescent axonal tracers were used to investigate the connections of medial agranular cortex (frontal area 2, Fr2) in male prairie voles. The rostral and caudal portions of Fr2 (rFr2 and cFr2) have distinct but partially overlapping patterns of connections. Thalamic labeling after cFr2 injections was present in anteromedial nucleus (AM), ventrolateral nucleus (VL), lateral segment, mediodorsal nucleus (MDl), centrolateral nucleus (CL), ventromedial nucleus (VM), posterior nucleus (Po) and lateral posterior nucleus (LP). A band of labeled cells involving CL, central medial nucleus (CM) and rhomboid nucleus (Rh) formed a halo around the periphery of submedial (gelatinosus) nucleus (Sm). Within cFr2 there is a rostrocaudal gradient whereby projections from VL and MDl become progressively sparser caudally, whereas those from LP and Po become denser. Rostral Fr2 receives afferents from a similar group of thalamic nuclei, but has denser innervation from VL and MDl, lacks afferents from LP, and receives less input from nuclei around the periphery of Sm. Caudal Fr2 has extensive cortical connections including orbital cortex, rostral Fr2, Fr1, caudal parietal area 1 (Par1), parietal area 2 (Par2), and posterior parietal, retrosplenial and visual areas. Rostral Fr2 has similar connections with areas Fr1, Par1 and Par2; orbital connections focused in ventrolateral orbital cortex (VLO); connections with caudal Fr2; greatly reduced connections with posterior parietal cortex and the visual areas; and no connections with retrosplenial cortex. The axons linking rFr2 and cFr2 with each other and with other cortical areas travel predominately in the deep gray matter of layers VI and VII rather than in the white matter. Projections to the dorsal striatum from rFr2 are widespread in the head of the caudate, become progressively restricted to a dorsocentral focus more caudally, and disappear by the level of the anterior commissure. The projections from cFr2 are largely restricted to a focal dorsocentral region of the striatum and to the dorsolateral margin of the caudatoputamen. In comparison to area Fr2, the laterally adjacent area Fr1 has thalamic and cortical connections which are markedly restricted. Area Fr1 receives thalamic input from nuclei VL, anteroventral nucleus (AV), CL and Po, but none from mediodorsal nucleus (MD) or LP, and its input from VM is reduced. Cortical afferents to Fr1 originate from areas Fr2, caudal Par1 and Par2. Medial agranular cortex of prairie voles has a pattern of connections largely similar to that seen in rats, suggesting that area Fr2 in prairie voles is part of a cortical network that may mediate complex behaviors involving spatial orientation. Received: 20 May 1998 / Accepted: 14 October 1998     

The distribution of the projection from the parataenial nucleus of the thalamus to the nucleus accumbens in the rat: An autoradiographic study

Summary In this study the intrastriatal distribution of afferents arising from the parataenial nucleus of the thalamus was investigated. Tritiated leucine and proline injected into the parataenial nucleus was found to densely label the entire anterior-posterior extent of the medial nucleus accumbens. The projection was for the most part limited to this striatal subregion, although some moderate labelling was found along the medial wall of the anterior caudateputamen. The terminal labelling within accumbens was characterized by a distinct patchiness. Other efferent connections of the parataenial nucleus observed in this study include the thalamic reticular nucleus, the basolateral and central nuclei of the amygdala, the septum, the medial frontal cortex, the entorhinal cortex and subiculum.This projection is distributed to the limbic afferented sector of striatum, and there is a nearly complete overlap between the parataenial afferents and those coming from hippocampus. The present findings suggest that the parataenial nucleus is an important thalamic link between limbic and striatal processing.Abbreviations A nucleus accumbens - AC anterior commissure - abl basolateral nucleus of amygdala - abm basomedial nucleus of amygdala - ac central nucleus of amygdala - am medial nucleus of amygdala - al lateral nucleus of amygdala - bst bed nucleus of Stria terminalis - CAI internal capsule - CC crus cerebri - CP caudate-putamen - F fornix - FCX frontal cortex - FH fimbria hippocampi - fr fasciculus retroflexus - GP globus pallidus - IL infralimbic cortex - lh lateral habenula - MD mediodorsal nucleus - MFH medial forebrain bundle - MO medial orbital cortex - nb nucleus basalis - ot olfactory tubercle - Pt parataenial nucleus - PV paraventricular nucleus - sl lateral septum - SM Stria medullaris - ST Stria terminalis - tam anterior medial nucleus - tav anterior ventral nucleus - tol nucleus of lateral olfactory tract - tr reticular nucleus - tv ventral nucleus - VL lateral ventricle - V_III third ventricle Presented in part at the Sixth European Neuroscience Congress, Malaga, Spain, 1982New address: Psychobiologie des Comportements Adaptatifs, INSERM U. 259, Rue Camille Saint Saens, Domaine de Carreire, F-33077 Bordeaux Cedex, France     

The termination of the spinothalamic tract in the cat an experimental study with silver impregnation methods

Summary The termination of the spinothalamic tract (STT) in the cat has been studied light microscopically in Fink-Heimer and Nauta impregnated sections. Following lesions of the STT at various rostrocaudal levels of the spinal cord the degenerating fibres in the thalamus and subthalamus were mapped, mainly in transverse sections. The cervicothalamic tract was not injured by the lesions.The spinothalamic fibres enter the diencephalon through the mesencephalic reticular formation and terminate in the following regions: the medial portion of the magnocellular part of the medial geniculate body (MGmc), the ventrolateral portion of the medial part of the posterior nuclear complex (PO_m), the caudolateral and medial parts of the zona incerta (ZI), the nucleus centralis medialis (CeM), the nucleus parafascicularis (Pf), the lateral part of the nucleus centralis lateralis (CL), the medial and rostrolateral parts of the nucleus ventralis lateralis (VL). To reach these regions the fibres pass through the nucleus centrum medianum (CM), the nucleus subparafascicularis (SPf) and the nucleus paracentralis (Pc). The fibres that terminate in the VL pass through Forel's field H₁ and the external medullated lamina (EML). Conclusive results were not obtained concerning a termination in the CM. The spinothalamic fibres do not pass through nor terminate in the nucleus ventralis posterolateralis (VPL) and the nucleus reticularis (R). The VPL, defined as that portion of the ventral thalamus that receives terminal fibres from the dorsal column nuclei, has been found to extend rostrally only as far as Horsley-Clarke level anterior 10.5. The results strongly support the view that all the spinothalamic fibres terminate ipsilateral to their course in the ventral quadrant of the spinal cord. No signs of a somatotopical organization of the termination of the STT were found.List of Abbreviations Cd nucleus caudatus - CeM nucleus centralis medialis - CG circumaqueductal gray substance - CL nucleus centralis lateralis thalami - CM nucleus centrum medianum thalami - CP commissura posterior - CTT cervicothalamic tract - EML external medullated lamina - H₁ Forel's field H₁ - HP tractus habenulopeduncularis - LCN nucleus cervicalis lateralis - LG corpus geniculatum laterale - LP nucleus lateralis posterior thalami - MD nucleus medialis dorsalis thalami - MG corpus geniculatum mediale - MGmc corpus geniculatum mediale, pars magnocellularis - MGp corpus geniculatum mediale, pars principalis - ML medial lemniscus - MRF mesencephalic reticular formation - OT optic tract - Pc nucleus paracentralis thalami - Pf nucleus parafascicularis thalami - PO posterior group of thalamic nuclei - PO₁ lateral part of PO - PO_m medial part of PO - R nucleus reticularis thalami - SG nucleus suprageniculatus - STT spinothalamic tract - VA nucleus ventralis anterior - VL nucleus ventralis lateralis thalami - VM nucleus ventralis medialis thalami - VPI nucleus ventralis posterior inferior - VPL nucleus ventralis posterior lateralis thalami (VPL₁ + VPL_m) - VPL₁ lateral part of VPL - VPL_m medial part of VPL - VPM nucleus ventralis posterior medialis thalami - VPMpc parvocellular part of VPM - ZI zona incerta     

Projections to the rostral reticular thalamic nucleus in the rat

Summary Afferent pathways to the rostral reticular thalamic nucleus (Rt) in the rat were studied using anterograde and retrograde lectin tracing techniques, with sensitive immunocytochemical methods. The analysis was carried out to further investigate previously described subregions of the reticular thalamic nucleus, which are related to subdivisions of the dorsal thalamus, in the paraventricular and midline nuclei and three segments of the mediodorsal thalamic nucleus. Cortical inputs to the rostral reticular nucleus were found from lamina VI of cingulate, orbital and infralimbic cortex. These projected with a clear topography to lateral, intermediate and medial reticular nucleus respectively. Thalamic inputs were found from lateral and central segments of the mediodorsal nucleus to the lateral and intermediate rostral reticular nucleus respectively and heavy paraventricular thalamic inputs were found to the medial reticular nucleus. In the basal forebrain, afferents were found from the vertical and horizontal limbs of the diagonal band, substantia innominata, ventral pallidum and medial globus pallidus. Brainstem projections were identified from ventrolateral periaqueductal grey and adjacent sites in the mesencephalic reticular formation, laterodorsal tegmental nucleus, pedunculopontine nucleus, medial pretectum and ventral tegmental area. The results suggest a general similarity in the organisation of some brainstem Rt afferents in rat and cat, but also show previously unsuspected inputs. Furthermore, there appear to be at least two functional subdivisions of rostral Rt which is reflected by their connections with cortex and thalamus. The studies also extend recent findings that the ventral striatum, via inputs from the paraventricular thalamic nucleus, is included in the circuitry of the rostral Rt, providing further evidence that basal ganglia may function in concert with Rt. Evidence is also outlined with regard to the possibility that rostral Rt plays a significant role in visuomotor functions.Abbreviations ac anterior commissure - aca anterior commissure, anterior - Acb accumbens nucleus - AI agranular insular cortex - AM anteromedial thalamic nucleus - AV anteroventral thalamic nucleus - BST bed nucleus of stria terminalis - Cg cingulate cortex - CG central gray - CL centrolateral thalamic nucleus - CM central medial thalamic nucleus - CPu caudate putamen - DR dorsal raphe nucleus - DTg dorsal tegmental nucleus - EP entopeduncular nucleus - f fornix - Fr2 Frontal cortex, area 2 - G gelatinosus thalamic nucleus - GP globus pallidus - Hb habenula - HDB horizontal limb of diagonal band - IAM interanterodorsal thalamic nucleus - ic internal capsule - INC interstitial nucleus of Cajal - IF interfascicular nucleus - IL infralimbic cortex - IP interpeduncular nucleus - LC locus coeruleus - LDTg laterodorsal tegmental nucleus - LH lateral hypothalamus - LHb lateral habenular nucleus - ll lateral lemniscus - LO lateral orbital cortex - LPB lateral parabrachial nucleus - MD mediodorsal thalamic nucleus - MDL mediodorsal thalamic nucleus, lateral segment - Me5 mesencephalic trigeminal nucleus - MHb medial habenular nucleus - mlf medial longitudinal fasciculus - MnR median raphe nucleus - MO medial orbital cortex - mt mammillothalamic tract - OPT olivary pretectal nucleus - pc posterior commissure - PC paracentral thalamic nucleus - PF parafascicular thalamic nucleus - PPTg pedunculopontine tegmental nucleus - PrC precommissural nucleus - PT paratenial thalamic nucleus - PV paraventricular thalamic nucleus - PVA paraventricular thalamic nucleus, anterior - R red nucleus - Re reuniens thalamic nucleus - RRF retrorubral field - Rt reticular thalamic nucleus - Scp superior cerebellar peduncle - SI substantia innominata - sm stria medullaris - SNR substantia nigra, reticular - st stria terminalis - TT tenia tecta - VL ventrolateral thalamic nucleus - VO ventral orbital cortex - VP ventral pallidum - VPL ventral posterolateral thalamic nucleus - VTA ventral tegmental area - 3 oculomotor nucleus - 3V 3rd ventricle - 4 trochlear nucleus     

The thalamo-caudate versus thalamo-cortical projections as studied in the cat with fluorescent retrograde double labeling

Summary The distribution of thalamic cells projecting to the head of the caudate and their interrelations with thalamo-cortical cells were studied in the cat with different combinations of fluorescent tracers. Injections in the head of the caudate were combined with the injections in the pericruciate, proreal, suprasylvian, anterior cingulate, occipital and ectosylvian cortices. The following results were obtained: (i) Injections in the head of the caudate resulted in retrograde labeling of thalamic cells medially and laterally to the anteromedial (AM) nucleus, and in the medioventral part of the ventral anterior (VA) nucleus. Further, labeled cells were distributed throughout the anterior intralaminar central medial (CeM), paracentral (Pc) and central lateral (CL) nuclei, and the posterior intralaminar center median-parafascicular complex (CM-Pf). Labeled cells were mainly grouped in the mediodorsal parts of the anterior intralaminar nuclei; they were also found in the more dorsal part of the mediodorsal (MD) nucleus, ventral to the thalamic paraventricular (Pv) nucleus and to the habenular complex, (ii) Thalamo-cortical and thalamo-caudate cells overlapped in the medial part of the VA; in the anterior intralaminar nuclei they were either intermingled or were distributed in separate clusters or longitudinal bands. The two cell populations also overlapped in the posterior intralaminar complex. The greatest overlap occurred with the thalamic cell population projecting to the pericruciate cortex. (iii) Thalamic cells bifurcating to the head of the caudate and to the pericruciate cortex were found lateral to the AM, within the VA, and throughout the anterior intralaminar nuclei, especially in the CeM and in the posterior part of the CL; a few branched cells were also found in the CM. Thalamic cells bifurcating to caudate and anterior suprasylvian cortex were also found in the VA. Very few cells (scattered in the anterior thalamus lateral to the AM, as well as in the CeM, Pc and CL) were found to bifurcate to the head of the caudate and the other cortical fields here examined.Supported in part by grants CNR 80.00515.04, 81.00283.04     

Acetylcholinesterase histochemistry in the macaque thalamus reveals territories selectively connected to frontal, parietal and temporal association cortices

The patterns of histochemical staining for acetylcholinesterase (AChE) activity in the macaque thalamus were analyzed and compared with the distribution of cells and terminals labeled from injections of axonal tracers in the dorsolateral and orbital prefrontal cortex, in area 7a of the posterior parietal cortex and in the polysensory cortex of the superior temporal sulcus. AChE histochemistry is very useful in delineating the thalamic nuclei connected with the association cortex and in uncovering thalamic subdivisions that are barely evident on cytoarchitectonic grounds. Moreover, AChE activity reveals previously unrecognized heterogeneities within several thalamic nuclei, like the ventral anterior (VA), where a new ventromedial subdivision (VAvm) is described, the medial pulvinar (PuIM) or the mediodorsal nucleus (MD). In this nucleus three distinct chemical domains are present: the medial, ventral and lateral sectors characterized by low, moderate and high AChE activities, respectively. The staining pattern of the lateral sector is markedly heterogeneous with patches of intense AChE activity surrounded by a moderately stained matrix. The MD medial sector is connected with the orbitofrontal cortex, whereas the AChE-rich patches in the lateral sector are selectively connected with the dorsolateral prefrontal, parietal and temporal association cortices. In the PulM, a dorsomedial AChE-rich patch is selectively connected with the orbitofrontal cortex, whereas the surrounding territory, which shows moderate AChE activity, is preferentially connected with the parietal and temporal cortices. Chemically specific domains in the anterior, ventral anterior, midline, and intralaminar thalamic nuclei are also connected with the examined association cortices. These findings indicate that the topographic patterns of the thalamo-cortical connections of primate association areas conform to the chemical architecture of the thalamus. This implies that because each cortical area is connected to a particular set of thalamic regions, the influence of the thalamus on cortical function is exclusive for each area, highly diverse among the various association areas, and subject to a wide range of modulation at the thalamic level.     

Topography and receptive field organization of the body midline representation in the ventrobasal complex of the cat

Summary The topography and receptive field (RF) organization of neurones in the trunk zone of the thalamic ventrobasal complex (VB) projecting to the homologous zone of the ipsilateral first somatosensory area (SI) were studied in the cat by performing experiments of retrograde neuronal tracing and microelectrode recording. Punctate cortical injections of small amounts of either horseradish peroxidase or fluorescent tracers (Evans Blue, Nuclear Yellow and Fast Blue) retrogradely labelled cell aggregates lying in the dorsal half of a VB region interposed between subnucleus VPL1 and VPLm. Aggregates of labelled cells were narrow in dorsoventral and mediolateral extent and elongated rostrocaudally. The distribution of VB cells projecting to the cortical subareas representing the dorsal midline, lateral trunk and ventral midline of the body in area SI, was established by injecting a different fluorescent marker into a physiologically defined site in each subarea. These injections resulted in labelling of three different cell aggregates located in topographically distinct regions of the VB trunk zone. Each aggregate of labelled cells only projected to one cortical subarea.Microelectrode analysis of cell populations of the VB trunk zone showed that neurones lying in regions projecting to dorsal and ventral midline zones of area SI had bilateral RFs, straddling the dorsal and the ventral midline of the body respectively. Neurones lying in the region projecting to the lateral trunk representation of area SI had contralateral RFs located on the lateral surface of the trunk.The results suggest that the detailed topography of the trunk map in the area SI and the bilaterality of the cortical representation of the body midlines, described in previous experiments, is imposed by the thalamocortical input from the VB.Abbreviations Ans Ansate sulcus - AV Nucleus anterior ventralis - Cd Nucleus caudatus - Ci Capsula interna - CL Nucleus centralis lateralis - CM Nucleus centrum medianum - Cru Cruciate sulcus - EB Evans blue - Ecsa Anterior ectosylvian sulcus - FB Fast blue - GL Nucleus geniculatus lateralis - Hb Nucleus habenularis - HRP Horseradish peroxidase - Lat Lateralis sulcus - LD Nucleus lateralis dorsalis - LP Nucleus lateralis posterior - MD Nucleus medialis dorsalis - NY Nuclear yellow - PC Nucleus paracentralis - Ped Pedunculus cerebri - PO Posterior thalamic group - POl Lateral division of the posterior thalamic group - POm Medial division of the posterior thalamic group - Pul Pulvinar - Psg Posterior sygmoid gyrus - R Nucleus reticularis thalami - Sups Suprasylvian sulcus - TO Tractus opticus - VA Nucleus ventralis anterior - VB Ventrobasal complex - VL Nucleus ventralis lateralis - VM Nucleus ventralis medialis - VPL1 Nucleus ventralis posterolateralis, lateral division - VPLm Nucleus ventralis posterolateralis, medial division Supported in part by funds granted by Italian Ministero della Pubblica Istruzione     

The vestibulothalamic projections in the cat studied by retrograde axonal transport of horseradish peroxidase

Summary Horseradish peroxidase (HRP) was injected or iontophoretically ejected in various thalamic nuclei in 63 adult cats. In 11 other animals HRP was deposited outside the thalamic territory. The number and distribution of labelled cells within the vestibular nuclear complex (VC) were mapped in each case. To a varying degree all subgroups of VC appear to contribute to the vestibulothalamic projections. Such fibres are distributed to several thalamic areas. From the present investigation it appears that generally speaking, there exist three distinct vestibulothalamic pathways with regard to origin as well as to site of termination of the fibres. One projection appears to originate mainly in caudal parts of the medial (M) and descending (D) vestibular nuclei and in cell group z. This pathway terminates chiefly in the contralateral medial part of the posterior nucleus of the thalamus (POm) including the magnocellular part of the medial geniculate body (Mgmc), the ventrobasal complex (VB) and the area of the ventral lateral nucleus (VL) bordering on VB. A second projection originates mainly in the superior vestibular nucleus (S) and in cell group y and terminates mainly in the contralateral nucleus centralis lateralis (CL) and the adjoining nucleus paracentralis (Pc). A third, more modest, pathway originates chiefly in the middle M and D, with a minor contribution from S and cell group y, and terminates in the contralateral ventral nucleus of the lateral geniculate body (GLV). There is some degree of overlap between the origin of these three vestibulothalamic pathways.Abbreviations B.c. brachium conjunctivum - CeM nucleus centralis medialis thalami - CL nucleus centralis lateralis thalami - CM nucleus centrum medianum - D nucleus vestibularis descendons - f cell group f - g cell group g - GLD corpus geniculatum laterale dorsalis - GLV corpus geniculatum laterale ventralis - i.e. nucleus intercalatus - L nucleus vestibularis lateralis - LD nucleus lateralis dorsalis thalami - LIM lamina medullaris interna - Lim nucleus limitans - LP nucleus lateralis posterior thalami - M nucleus vestibularis medialis - MD nucleus medialis dorsalis thalami - MGmc corpus geniculatum mediale, pars magnocellularis - MGp corpus geniculatum mediale, pars principalis - N.cu.e. nucleus cuneatus externus - N.f.c. nucleus fasciculi cuneati - N.mes. V nucleus mesencephalicus nervi trigemini - NR nucleus ruber - N.tr.s. nucleus tractus solitarius - N. VII nervus facialis - N. VIII nervus statoacusticus - PC pedunculus cerebri - Pc nucleus paracentralis thalami - Pf nucleus parafascicularis - p.h. nucleus prepositus hypoglossi - PO posterior thalamic group - PO1 lateral part of PO - POm medial part of PO - Prt nucleus pretectalis - Pul pulvinar - R nucleus reticularis thalami - S nucleus vestibularis superior - Sg nucleus suprageniculatus - SN substantia nigra - Sv nucleus supravestibularis - Tr.s. tractus solitarius - VA nucleus ventralis anterior thalami - VL nucleus ventralis lateralis thalami - VPL nucleus ventralis posterior lateralis - VPL1 lateral part of VPL - VPLm medial part of VPL - VPM nucleus ventralis posterior medialis - x cell group x - y cell group y - z cell group z - V nucleus motorius nerve trigemini - X nucleus dorsalis nerve vagi - XII nucleus nervi hypoglossi     

The mode of nigro-thalamic transmission investigated with intracellular recording in the cat

Postsynaptic potentials evoked by stimulating the substantia nigra (SN) were recorded intracellularly from ipsilateral ventral medial (VM), ventral lateral (VL), and ventral anterior (VA) nuclei of the thalamus in cats anesthetized with sodium pentobarbital. SN stimulation evoked inhibitory postsynaptic potentials (IPSP) at a short latency in VM neurons (mean 1.68 ms, SD 0.23, n = 59). The IPSP were produced monosynaptically because linear regression analysis of latency vs. conduction distance between stimulating and recording sites indicated a synaptic delay of less than 0.6 ms. Conduction velocity for these fibers was calculated to be 4.48 m/s. The spots from which IPSP were produced with the lowest threshold were determined for each of 38 VM neurons. IPSP origins thus determined were distributed in the pars reticulata of the SN (SNr) and in the area where nigro-thalamic fibers run. Neurons which received IPSP from the SNr were distributed in the VM nucleus, ventromedial to the VL nucleus, where fibers from the contralateral brachium conjunctivum terminate. Convergence of nigral and cerebellar impulses was not observed in thalamic neurons sampled in this study. Stimulation of the entopeduncular nucleus (ENT) also produced monosynaptic IPSP in VL-VA neurons. The SNr-related cell group was located ventromedially and caudally to the ENT-related cell group. No convergence of nigral and pallidal influences was observed within thalamic neurons.     

Differential projection from the motor and limbic cortical regions to the mediodorsal thalamic nucleus in the dog

The cortical afferents to the mediodorsal thalamic nucleus in the dog were studied by using horseradish peroxidase. Small injections allowed to establish two specific projection zones connected separately with the lateral and medial segments of the nucleus. The lateral segment received the major projection from the dorsal half of the hemisphere. It included premotor and part of the motor cortices in the anterior sigmoid gyrus and precruciate areas as well as the presylvian cortex. The medial segment of the nucleus was innervated by the limbic areas of the ventral half of the hemisphere. These areas included the medioventrally located genual, subcallosal and piriform cortices, as well as the cortex of the ventral bank of the anterior rhinal sulcus and the caudal part of the orbital gyrus. The cortical fields situated between these two main cortical zones, both on the lateral and medial surfaces (rhinal and sylvian sulci and anterior cingular gyrus, respectively) sent projections to both medial and lateral segments of the nucleus. These results indicate that in the mediodorsal thalamic nucleus may take place the integration of information from two functionally defined systems, the motor and limbic ones.     

Direct projections of the hypothalamic nuclei to the thalamic mediodorsal nucleus in the cat

Direct projections of the hypothalamic nuclei to the thalamic mediodorsal nucleus (MD) were studied using retrograde and anterograde transport of horseradish peroxidase (HRP) and wheat germ agglutinin (WGA)-HRP. HRP and WGA-HRP were injected into the MD, thalamic paraventricular, lateral habenular and hypothalamic nuclei. The results indicate that the MD, particularly its medial part, receives a moderate amount of hypothalamic afferents, and that most of these afferents originate in the medial part of the lateral hypothalamic nucleus at anterior levels, while a limited number are derived from the dorsal, dorsomedial, ventromedial and anterior hypothalamic and lateral preoptic nuclei.