An investigation of collateral projections of the dorsal lateral geniculate nucleus and other subcortical structures to cortical areas V1 and V4 in the macaque monkey: A double label retrograde tracer study

Summary Previous anterograde studies in the macaque monkey have shown that, in addition to the projection to striate cortex (V1), the dorsal lateral geniculate nucleus (DLG) has a sparse, horizontally segregated projection to layers IV and V of prestriate cortex (V4). However, the distribution and degree of axon collateralization of DLG cells which give rise to these projections are unknown. This study was designed to answer these questions. The DLG (along with the pulvinar and other subcortical regions) was examined for the presence of single- or doublelabeled cells after injections of two different (fluorescent or HRP) retrograde tracers into corresponding retinotopic points in visual cortical areas V1 and V4. In the DLG, it was found that cells projecting to V4, which reside in or near the tectorecipient interlaminar zones of the DLG, do not project to V1 and thus represent a separate population of cells. The organization of the macaque geniculo-prestriate projection thus seems quite different from that of carnivores. Both single- and double-labeled cells were found in other subcortical areas, e.g., single-labeled cells were found in the claustrum, hypothalamus and lateral pulvinar, and a double-labeled cell population was found in the inferior pulvinar.Abbreviations BSC Brachium of the superior colliculus - Cd Caudate nucleus - Cl Claustrum - DLG Dorsal lateral geniculate nucleus - GP Globus pallidus - Hce1 Medial hypothalamic cholinesterase group of Mesulam et al. (1983) - Hce2 Lateral hypothalamic cholinesterase group of Mesulam et al. (1983) - Hip Hippocampus - ot optic tract - PI Inferior pulvinar - PL Lateral pulvinar - PM Medial pulvinar - Put Putamen - Ret Reticular nucleus of thalamus - Thal Thalamus - v ventricle     

Network organization of the connectivity between parietal area 7, posterior cingulate cortex and medial pulvinar nucleus: a double fluorescent tracer study in monkey

Summary The topographic distribution of medial pulvinar neuronal populations projecting to area 7a and to posterior cingulate gyrus (area 23) was investigated with retrograde axonal transport of fluorescent dyes. In an initial stage, separate injections of fast blue and diamidino-yellow were placed in area 7a. Two segregated backfilled cell populations were observed in the cingulate gyrus revealing a topographic distribution of cortico-cortical connections. In a second stage, the two dyes were injected in area 7a and in the posterior cingulate gyrus. After injections in two cortical sub-areas previously shown to be in topographic correspondence, the two projecting populations observed in the medial pulvinar over-lapped extensively. In the case of injections in two regions not topographically correspondent, the two medial pulvinar populations were found to be segregated. These findings reveal that within both posterior parietal cortex and posterior cingulate gyrus, sub-areas which are in topographic correspondence receive projections from a common region of the medial pulvinar.     

Connections between the pulvinar complex and cytochrome oxidase-defined compartments in visual area V2 of macaque monkey

We examined the distribution of pulvinar afferents to visual area V2 of macaque monkey cerebral cortex in relation to the distribution of the metabolic enzyme cytochrome oxidase (CO). V2 contains three sets of stripelike subregions that are marked by differential staining for CO, and which have different corticocortical connections. The pulvinar provides the major subcortical input to V2, and this input is known to be patchy. We were interested to determine how the pattern of pulvinar afferents relates to the layout of the three stripelike compartments that characterize V2. We made large injections of WGA-HRP into the pulvinar (labelling both the inferior and lateral divisions) and mapped the resulting orthograde terminal and retrograde cell label within V2. We observed pulvinar terminal label mainly in lower layer 3 (at the layer 4 border), with light label in layer 1 as well; terminal label in layers 3–4 was distributed in discrete patches with faint bridges of light label between. Comparison with adjacent sections stained for CO or Cat-301 showed that pulvinar terminal zones aligned precisely with regions of increased CO staining, and targeted both thick (Cat-301⁺) and thin CO-rich stripes, avoiding the pale stripes (which aligned with the faint bridges of terminal label). Retrogradely labelled cells were found in layers 5A and 6, but the bulk of the feedback to pulvinar arose from layer 6 rather than layer 5 (unlike V1, where feedback to pulvinar arises primarily from layer 5B). These results show that the increased CO staining in certain subregions of V2 is closely correlated with the presence of thalamic terminals from the pulvinar. Although we cannot rule out the possibility that different sets of pulvinar neurons project to different CO compartments in V2, the presence of a prominent thalamic input shared by the thick and thin CO stripes (which receive different V1 afferents and make different feedforward projections to other visual cortical areas) could underlie the preferential intrinsic interconnections shown to exist between these V2 subregions and suggests another potential source of integration between the two cortical visual streams.     

A horseradish peroxidase-autoradiographic study of parietopulvinar connections in saimiri sciureus

Summary Descending connections from parietal cortex to pulvinar in squirrel monkey were investigated with the autoradiographic method. Somatosensory areas I (SI) and II (SII) were found to project to the oral (PuO) and medial (PuM) subdivisions of the pulvinar. Projections from the posterior parietal region were recorded in circumscribed areas of PuM and the lateral (PuL) and inferior (PuI) subdivisions of pulvinar. Retrograde labeling with horseradish peroxidase (HRP) demonstrated that rostral parietal cortex including the lateral cortex of SI and the rostral part of area 5 received reciprocal projections from PuO and rostral PuM. Injections of HRP into medial and lateral regions of SI also resulted in labeled cells in PuO and PuM. Within the limitations of the HRP technique, the latter results suggest a direct pathway from pulvinar to primary sensory cortex. The experimental results confirm the accepted view of projections from parieto-temporo-occipital association cortex to PuM, PuL and PuI. In addition, reciprocal connections of rostral parietal cortex with PuO and PuM were demonstrated.     

Visual thalamocortical projections in the flying fox: parallel pathways to striate and extrastriate areas

We studied thalamic projections to the visual cortex in flying foxes, animals that share neural features believed to resemble those present in the brains of early primates. Neurones labeled by injections of fluorescent tracers in striate and extrastriate cortices were charted relative to the architectural boundaries of thalamic nuclei. Three main findings are reported: First, there are parallel lateral geniculate nucleus (LGN) projections to striate and extrastriate cortices. Second, the pulvinar complex is expansive, and contains multiple subdivisions. Third, across the visual thalamus, the location of cells labeled after visual cortex injections changes systematically, with caudal visual areas receiving their strongest projections from the most lateral thalamic nuclei, and rostral areas receiving strong projections from medial nuclei. We identified three architectural layers in the LGN, and three subdivisions of the pulvinar complex. The outer LGN layer contained the largest cells, and had strong projections to the areas V1, V2 and V3. Neurones in the intermediate LGN layer were intermediate in size, and projected to V1 and, less densely, to V2. The layer nearest to the origin of the optic radiation contained the smallest cells, and projected not only to V1, V2 and V3, but also, weakly, to the occipitotemporal area (OT, which is similar to primate middle temporal area) and the occipitoparietal area (OP, a "third tier" area located near the dorsal midline). V1, V2 and V3 received strong projections from the lateral and intermediate subdivisions of the pulvinar complex, while OP and OT received their main thalamic input from the intermediate and medial subdivisions of the pulvinar complex. These results suggest parallels with the carnivore visual system, and indicate that the restriction of the projections of the large- and intermediate-sized LGN layers to V1, observed in present-day primates, evolved from a more generalized mammalian condition.     

Thalamic projections to visual and visuomotor areas (V6 and V6A) in the Rostral Bank of the parieto-occipital sulcus of the Macaque

The medial posterior parietal cortex of the primate brain includes different functional areas, which have been defined based on the functional properties, cyto- and myeloarchitectural criteria, and cortico-cortical connections. Here, we describe the thalamic projections to two of these areas (V6 and V6A), based on 14 retrograde neuronal tracer injections in 11 hemispheres of 9 Macaca fascicularis. The injections were placed either by direct visualisation or using electrophysiological guidance, and the location of injection sites was determined post mortem based on cyto- and myeloarchitectural criteria. We found that the majority of the thalamic afferents to the visual area V6 originate in subdivisions of the lateral and inferior pulvinar nuclei, with weaker inputs originating from the central densocellular, paracentral, lateral posterior, lateral geniculate, ventral anterior and mediodorsal nuclei. In contrast, injections in both the dorsal and ventral parts of the visuomotor area V6A revealed strong inputs from the lateral posterior and medial pulvinar nuclei, as well as smaller inputs from the ventrolateral complex and from the central densocellular, paracentral, and mediodorsal nuclei. These projection patterns are in line with the functional properties of injected areas: “dorsal stream” extrastriate area V6 receives information from visuotopically organised subdivisions of the thalamus; whereas visuomotor area V6A, which is involved in the sensory guidance of arm movement, receives its primary afferents from thalamic nuclei that provide high-order somatic and visual input.     

Topographical and topological organization of the thalamocortical projection to the striate and prestriate cortex in the marmoset (Callithrix jacchus)

Summary In eleven hemispheres of nine marmoset monkeys (Callithrix jacchus), we have investigated the thalamo-cortical organization of the projections from the pulvinar to the striate and prestriate cortex. In each experiment, single or multiple injections of various retrograde fluorescent tracers were injected into adjacent regions or areas. In two experiments, horseradish peroxidase (HRP) was injected into the lateral geniculate nucleus (LGN) and the lateral pulvinar, respectively. The results show that the thalamo-cortical projection from LGN to striate cortex and from pulvinar to the prestriate cortex are similarly organized, but the geniculo-striate projection is more precise than the pulvinar-prestriate projection. The pulvinar-prestriate projection is topographically organized and preserves topological neighbourhood relations. Projection zones to the various visual areas are concentrically wrapped around each other. The projection zone to area 18 constitutes a central core region. It begins ventro-laterally in PuL where the pulvinar is in contact with the LGN. This contact zone we called the hilus region of the pulvinar. The area 18-projection zone stretches as a central cone into the posterior pulvinar through PuL and into PuM. It is surrounded by the projection zone to the posterior belt of area 19 and this in turn is surrounded by the projection zone to the anterior belt of area 19. The projection zones to area 19 are then surrounded medially and dorsally by zones projectiong to the temporal and parietal association cortex, respectively. The projection zone to area MT is located medio-ventrally in the posterior pulvinar (PuIP and surrounding nuclei) and coincides with a densely myelinated region. Area 17 also receives input from the pulvinar but probably predominantly in the region of the central visual field. The pulvinar zone projecting to area 17 is located ventrolaterally from the central core region projecting to area 18 and is contiguous laterally with the LGN. If the positions of the vertical and the horizontal meridian in the pulvinar correspond to those in the respective cortical projection zones, a second order visual field representation such as found in area 18, with the horizontal meridian split at an excentricity of about 7–10°, can also be recognized in the pulvinar.Abbreviations A Subcortical nuclei and subnuclei, cf. — Stephan et al. (1980) - AD Nucleus anterior dorsalis thalami - AV Nucleus anterior ventralis thalami - CeD Nucleus centralis dorsalis thalami - CeL Nucleus centralis lateralis thalami - CeMe Centrum medianum thalami - CoS Colliculus superior - FRPO Formatio reticularis pontis, pars oralis - GM Corpus geniculatum mediale - IBCI Nucleus interstitialis brachii colliculi inferioris - LGN Corpus geniculatum laterale dorsale - vLGN Corpus geniculatum laterale ventrale - LD Nucleus lateralis dorsalis thalami - LI Nucleus limitans thalami - LP Nucleus lateralis posterior thalami - MD Nucleus medialis dorsalis thalami - OL Nucleus olivaris superior lateralis - OM Nucleus olivaris superior medialis - Pbg Nucleus parabigeminalis - Pul Pulvinar inferior; PulP Pulvinar inferior posterior - PuL Pulvinar lateralis - PuM Pulvinar medialis - PuO Pulvinar oralis - RT Nucleus reticularis thalami - Sg Nucleus suprageniculatus - VA Nucleus ventralis anterior thalami - VL Nucleus ventralis lateralis thalami - VPL Nucleus ventralis posterior lateralis thalami - VPM Nucleus ventralis posterior medialis thalami - IV Nucleus nervi trochlearis - B Cortical areas and subareas, (after Spatz 1977a; Spatz et al. 1987 Allman and Kaas 1975): - 17 Area striata (V I) - 18 Area 18 (V II) - 19DI Area 19 dorso-intermediate - 19DL Area 19 dorso-lateral - 19DM Area 19 dorso-medial - 19M Area 19 medial - 19V Area 19 ventral - MT Middle temporal area     

Parietal inputs to dorsal versus ventral premotor areas in the macaque monkey: evidence for largely segregated visuomotor pathways

The lateral premotor cortex plays a crucial role in visually guided limb movements. Visual information may reach this cortical region from the parietal cortex, the highest stage in the dorsal visual stream. Anatomical studies indicate that the parietal projections to the dorsal (PMd) and ventral (PMv) premotor areas arise from separate parietal regions, supporting the notion of parallel visuomotor pathways. We tested the degree of segregation of these pathways by injecting retrograde tracers into PMd and PMv in the same monkeys, under physiological control. Eleven injections were made in four animals, and the analysis of retrograde labelling revealed that parietal cells projecting to PMd and those projecting to PMv are largely segregated. The strongest projections to PMd arise from the superior parietal lobule, including the medial intraparietal area (MIP), PEc and PGm, and the parieto-occipital area. These areas were devoid of labelling following injections into PMv, which receives its major projections from the anterior intraparietal area (AIP), area PEip, the anterior portion of the inferior parietal gyrus (area 7b), and the somatosensory areas. In addition to their strong projections to PMv, areas 7b and PEip send minor projections to PMd as well. Additional projections to PMd arise from the ventral intraparietal area and the inferior parietal lobule. The present findings are direct anatomical evidence for largely segregated visuomotor pathways linking parietal cortex with PMd and PMv.     

Retinotopy and color sensitivity in human visual cortical area V8

Prior studies suggest the presence of a color-selective area in the inferior occipital-temporal region of human visual cortex. It has been proposed that this human area is homologous to macaque area V4, which is arguably color selective, but this has never been tested directly. To test this model, we compared the location of the human color-selective region to the retinotopic area boundaries in the same subjects, using functional magnetic resonance imaging (fMRI), cortical flattening and retinotopic mapping techniques. The human color-selective region did not match the location of area V4 (neither its dorsal nor ventral subdivisions), as extrapolated from macaque maps. Instead this region coincides with a new retinotopic area that we call 'V8', which includes a distinct representation of the fovea and both upper and lower visual fields. We also tested the response to stimuli that produce color afterimages and found that these stimuli, like real colors, caused preferential activation of V8 but not V4.     

Neocortical connections in fetal cats

The major extrinsic projections to and from visual and auditory areas of cerebral cortex were examined in fetal cats between 46 and 60 days of gestation (E46-E60) using axonal transport of horseradish peroxidase either alone or in combination with tritiated proline. Projections to visual cortex from the dorsal lateral geniculate nucleus and lateral-posterior/pulvinar complex exist by E46, and those from the contralateral hemisphere, claustrum, putamen, and central lateral nucleus of the thalamus are present by E54-E56. In addition, cells in the medial geniculate nucleus project to auditory cortex by E55. At E54-E56 efferent cortical projections reach the contralateral hemisphere, claustrum, putamen, lateral-posterior/pulvinar complex and reticular nucleus of the thalamus. Cells in visual cortex also project to the dorsal and ventral lateral geniculate nuclei, pretectum, superior colliculus and pontine nuclei, and cells in auditory cortex project to the medial geniculate nucleus. Except for interhemispheric projections, all pathways demonstrated are ipsilateral, and projections linking cerebral cortex with claustrum, dorsal lateral geniculate nucleus and lateral-posterior/pulvinar complex are reciprocal. The reciprocal projections formed with the dorsal lateral geniculate nucleus, lateral-posterior/pulvinar complex and the claustrum show a greater degree of topological organization compared to the projections formed with the contralateral hemisphere and superior colliculus, which show little or no topological order. Therefore, the results of the present study show that the major extrinsic projections of the cat's visual and auditory cortical areas with subcortical structures are present by the eighth week of gestation, and that the origins and terminations of many of these projections are arranged topologically.     

Afferent connections of the prelunate visual association cortex (areas V4 and DP)

Summary The afferent and efferent connections of the prelunate visual association area V4 of macaque monkeys were investigated by means of the horseradish peroxidase (HRP) method. The specific thalamic afferents from the dorsolateral segment of the medial pulvinar and the lateral segment of the inferior pulvinar were topographically organized. A band of cells was labelled in the intralaminar nuclei (nucl. centr. med. and lat., reaching into LD and the most dorsal part of VL), and a few cells in the interlaminar layers of the lateral geniculate body. Other diencephalic afferents included the claustrum, the nucleus basalis Meynert and the pars compacta of the substantia nigra. Ipsilateral cortical areas which projected into V4 included area 18 (V2), the inferior parietal cortex, the anterior and posterior parts of the superior temporal sulcus, the frontal eye fields and the temporo-basal association cortex on the lateral half of the parahippocampal gyrus and around the occipito-temporal sulcus. In the contralateral cortex, discontinuous regions in areas V4 and V5 on the prelunate gyrus and some cells at the 17/18-border were labelled. All regions in which labelled cells were found and, in addition a restricted region in the dorsal cap of the head and the tail of the caudate nucleus showed fibre and terminal labelling. In addition mesencephalic afferents and efferents were identified but not investigated in detail. An attempt to estimate the quantitative contribution of the various afferent systems to the prelunate cortex was made by counting the labelled cells in the different areas. The afferent and efferent organization of the prelunate visual association area indicates that it is incorporated in a network of cortical and subcortical regions involved in various aspects of visual behavior.     

Distribution of neurons in the main cuneate nucleus projecting to the inferior olive in the cat. Evidence that they differ from those directly projecting to the cerebellum

The distribution in the main cuneate nucleus of cells projecting to the inferior olive and the intermediate zone of the cerebellar anterior lobe were compared by means of double retrograde labeling methods in the cat. The tracer combinations were either Fast Blue and Diamidino Yellow Dihydrochloride; or horseradish peroxidase conjugated to wheat germ agglutinin and Diamidino Yellow Dihydrochloride. Neurons in the caudal, middle and rostral subdivisions of the main cuneate nucleus project to the inferior olive. Differences exist, however, in its number and location along the rostrocaudal extent of the nucleus. Cells projecting to the inferior olive predominate in the caudal and middle subdivisions, where they concentrate ventrally. No cells in the "clusters region" project to the inferior olive. Main cuneate nucleus neurons projecting to the cerebellum concentrate rostral to the obex, bordering the external cuneate nucleus and partially intermixing with the rostrally located cells projecting to the inferior olive. However, no double-labeled cells were found. The results indicate that the main cuneate nucleus projections to the inferior olive and cerebellar anterior lobe originate from different populations of neurons with high specific locations within the nucleus. This finding is in agreement with previous studies suggesting that each of the main cuneate nucleus targets receives its input from a distinct population of neurons within the nucleus.     

Brain stem projections to the pulvinar-lateralis posterior complex of the cat

Summary The present experiments were undertaken to define the areas of projection of pretectum and superior colliculus to the pulvinar and n. lateralis posterior, respectively, and to define other brain stem structures projecting to these thalamic nuclei in cats. For this purpose the technique of retrograde transport of horseradish peroxidase (HRP) has been used.After injection of the enzyme in the pulvinar, neurons were labeled in all subdivisions of the pretectal area. The majority of the labeled cells were located in the n. pretectalis posterior and n. tractus opticus although cells filled with HRP were present also in the n. pretectalis anterior pars compacta and area pretectalis medialis. Neurons projecting to the pulvinar were also found in the periaqueductal gray, reticular formation and locus coeruleus.When HRP was injected in the n. lateralis posterior, labeled neurons were present in the II and III subdivisions of the second layer of the superior colliculus. The location of these cells shifted from medial to lateral as the injections were shifted from posterior to anterior within the lateralis posterior. Neurons projecting to this nucleus were also present in the intermediate layers of the superior colliculus, lateral hypothalamus and parabigeminal nucleus.The possible role of the pretectal area and superior colliculus in mediating somesthetic input to the pulvinar and lateralis posterior, respectively, and the role of these structures in the control of ocular movements, are discussed.Abbreviations APM area pretectalis medialis - Cu nucleus cuneiformis - CS nucleus centralis superior - fr fasciculus retroflexus - Gp pontine gray - Hb nucleus habenulae - IC inferior colliculus - LC locus coeruleus - LGB lateral geniculate body - LP lateralis posterior - MGB medial geniculate body - nPA_c nucleus pretectalis anterior pars compacta - nPA_r nucleus pretectalis anterior pars reticularis - nPC nucleus posterior commissurae - nPP nucleus pretectalis posterior - nTO nucleus tractus opticus - PAG periaqueductal gray - PB nucleus parabigeminalis - Pi pulvinar inferior - PO nucleus posterior of the thalamus - Pul pulvinar - Pt pretectum - RF reticular formation - Rtp tegmental reticular nucleus - SC superior colliculus Supported by H. de Jur Foundation and USPHS Grant TWO 2718Present address: Max-Planck-Institut für biophysikalische Chemie, Postfach 968, D-3400 Göttingen, Federal Republic of Germany     

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.     

Corticothalamic connections of the superior temporal sulcus in rhesus monkeys

Summary The corticothalamic connections of the superior temporal sulcus (STS) were studied by means of the autoradiographic technique. The results indicate that corticothalamic connections of the STS in general reciprocate thalamocortical connections. The cortex of the upper bank of the STS-multimodal areas TPO and PGa-projects to four major thalamic targets: the pulvinar complex, the mediodorsal nucleus, the limitanssuprageniculate nucleus, as well as intralaminar nuclei. Within the pulvinar complex, the main projections of the upper bank of the STS are directed to the medial pulvinar (PM) nucleus. Rostral upper bank regions tend to project caudally and medially within the PM nucleus, caudal upper bank regions, more laterally and ventrally. The mid-portion of the upper bank tends to occupy the central sector of the PM nucleus. There are also relatively minor projections from upper bank regions to the lateral pulvinar (PL) and oral pulvinar (PO) nuclei. In contrast to the upper bank, the projections from the lower bank are directed primarily to the pulvinar complex, with only minor projections to intralaminar nuclei. The rostral portion of the lower bank projects mainly to caudal and medial regions of the PM nucleus, whereas the caudal lower bank projects predominantly to the lateral PM nucleus, and also to the PL, PO, and inferior pulvinar (PI) nuclei. The mid-portion of the lower bank projects mainly to central and lateral portions of the PM nucleus, and also to the PI and PL nuclei. The rostral depth of the STS projects mainly to the PM nucleus, with only minor connections to the PO, PI, and PL nuclei. The midportion of multimodal area TPO of the upper bank, areas TPO₂ and TPO₃, projects preferentially to the central sector of the PM nucleus. It is possible that this STS-thalamic connectivity has a role in behavior that is dependent upon more than one sensory modality.Abbreviations AM anterior medial nucleus - AS arcuate sulcus - AV anterior ventral nucleus - BSC brachium of the superior colliculus - Cd caudate nucleus - Cif nucleus centralis inferior - Cim nucleus centralis intermedialis - CL central lateral nucleus - CM centromedian nucleus - CM-Pf centromedian-parafascicular nucleus - Cs nucleus centralis superior - CS central sulcus - CSL nucleus centralis lateralis superior - GLd dorsal lateral geniculate nucleus - GM medial geniculate nucleus - Hb habenula - IOS inferior occipital sulcus - IPS intraparietal sulcus - LD lateral dorsal nucleus - LF lateral fissure - Li limitans nucleus - LP lateral posterior nucleus - LS lunate sulcus - MD mediodorsal nucleus - Pa paraventricular nucleus - Pen paracentral nucleus - Pf parafascicular nucleus - PI inferior pulvinar nucleus - PL lateral pulvinar nucleus - PM medial pulvinar nucleus - PO oral pulvinar nucleus - PS principal sulcus - Pt parataenial nucleus - R reticular nucleus - Re reuniens nucleus - SG suprageniculate nucleus - STN subthalamic nucleus - STS superior temporal sulcus - THI habenulo-interpeduncular tract - VLc nucleus ventralis lateralis, pars caudalis - VLm nucleus ventralis lateralis, pars medialis - VLo nucleus ventralis lateralis, pars oralis - VLps nucleus ventralis lateralis, pars postrema - VPI ventroposteroinferior nucleus - VPLc nucleus ventralis posterior lateralis, pars caudalis - VPLo nucleus ventralis posterior lateralis, pars oralis - VPM ventroposteromedial nucleus - VPMpc ventroposteromedial nucleus, parvocellular portion - X nucleus X     

Microstructural organizational patterns in the human corticostriatal system

The axons that project into the striatum are known to segregate according to macroscopic cortical systems; however, the within-region organization of these fibers has yet to be described in humans. We used in vivo fiber tractography, in neurologically healthy adults, to map white matter bundles that originate in different neocortical areas, navigate complex fiber crossings, and project into the striatum. As expected, these fibers were generally segregated according to cortical origin. Within a subset of pathways, a patched pattern of inputs was observed, consistent with previous ex vivo histological studies. In projections from the prefrontal cortex, we detected a topography in which fibers from rostral prefrontal areas projected mostly to rostral parts of the striatum and vice versa for inputs originating in caudal cortical areas. Importantly, within this prefrontal system there was also an asymmetry in the subset of divergent projections, with more fibers projecting in a posterior direction than anterior. This asymmetry of information projecting into the basal ganglia was predicted by previous network-level computational models. A rostral-caudal topography was also present at the local level in otherwise somatotopically organized fibers projecting from the motor cortex. This provides clear evidence that the longitudinal organization of input fields, observed at the macroscopic level across cortical systems, is also found at the microstructural scale at which information is segregated as it enters the human basal ganglia.     

Thalamocortical and the dual pattern of corticothalamic projections of the posterior parietal cortex in macaque monkeys

The corticothalamic projection includes a main, modulatory projection from cortical layer VI terminating with small endings whereas a less numerous, driving projection from layer V forms giant endings. Such dual pattern of corticothalamic projections is well established in rodents and cats for many cortical areas. In non-human primates (monkeys), it has been reported for the primary sensory cortices (A1, V1, S1), the motor and premotor cortical areas and, in the parietal lobe, also for area 7. The present study aimed first at refining the cytoarchitecture parcellation of area 5 into the sub-areas PE and PEa and, second, establishing whether area 5 also exhibits this dual pattern of corticothalamic projection and what is its precise topography. To this aim, the tracer biotinylated dextran amine (BDA) was injected in area PE in one monkey and in area PEa in a second monkey. Area PE sends a major projection terminating with small endings to the thalamic lateral posterior nucleus (LP), ventral posterior lateral nucleus (VPL), medial pulvinar (PuM) and, but fewer, to ventral lateral posterior nucleus, dorsal division (VLpd), central lateral nucleus (CL) and center median nucleus (CM), whereas giant endings formed restricted terminal fields in LP, VPL and PuM. For area PEa, the corticothalamic projection formed by small endings was found mainly in LP, VPL, anterior pulvinar (PuA), lateral pulvinar (PuL), PuM and, to a lesser extent, in ventral posterior inferior nucleus (VPI), CL, mediodorsal nucleus (MD) and CM. Giant endings originating from area PEa formed restricted terminal fields in LP, VPL, PuA, PuM, MD and PuL. Furthermore, the origin of the thalamocortical projections to areas PE and PEa was established, exhibiting clusters of neurons in the same thalamic nuclei as above, in other words predominantly in the caudal thalamus. Via the giant endings CT projection, areas PE and PEa may send feedforward, transthalamic projections to remote cortical areas in the parietal, temporal and frontal lobes contributing to polysensory and sensorimotor integration, relevant for visual guidance of reaching movements for instance.     

Development of banded afferent compartments in the inferior colliculus before onset of hearing in ferrets

Axonal projections from the lateral superior olivary nuclei (LSO), as well as from the dorsal cochlear nucleus (DCN) and dorsal nucleus of the lateral lemniscus (DNLL), converge in frequency-ordered layers in the central nucleus of the inferior colliculus (IC) where they distribute among different synaptic compartments. A carbocyanine dye, 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate (DiI), was used as a tracer to study the postnatal development of axonal projections in the ferret IC. The results indicated that projections from all three nuclei are present at birth, but are not segregated into bands. During the postnatal week between approximately postnatal days 4 and 12 (P4-P12), axons from LSO proliferate in IC, become more branched, and segregate into a series of bands composed of densely packed fibers and endings. LSO projections in these afferent bands course parallel to IC layers and are separated by intervening regions with few endings. A modest fit of a sine curve (R2>0.15) to the pattern of spacing of LSO projections in IC indicated that regularly spaced bands are forming by P7. Similarly, banded patterns of DCN and DNLL projections to IC have developed by the end of the first postnatal week. Thus, well before hearing onset in ferret (P28-30), three different afferent projections have segregated into banded compartments along layers in the central nucleus of the ferret IC. Possible mechanisms in circuit development are discussed.     

Projections from inferior temporal cortex to prefrontal cortex via the uncinate fascicle in rhesus monkeys

Summary In five rhesus monkeys (Macaca, mulatta) we used anterograde and retrograde tracing techniques to investigate the projection from the inferior temporal cortex (area TE) to the prefrontal cortex as well as the course of the projecting fibers. The results showed that TE projects to both the inferior convexity and orbital surface of prefrontal cortex and that these projections course almost exclusively via the uncinate fascicle. Transection of the uncinate fascicle deprives the prefrontal cortex of virtually all input from TE, but leaves intact inputs from prestriate and parietal visual areas as well as the amygdala. Such transection also leaves intact many projections from TE to targets other than the prefrontal cortex, including the amygdala, ventral putamen, tail of the caudate nucleus, and pulvinar.     

Effect of attentive fixation in macaque thalamus and cortex

Attentional modulation of neuronal responsiveness is common in many areas of visual cortex. We examined whether attentional modulation in the visual thalamus was quantitatively similar to that in cortex. Identical procedures and apparatus were used to compare attentional modulation of single neurons in seven different areas of the visual system: the lateral geniculate, three visual subdivisions of the pulvinar [inferior, lateral, dorsomedial part of lateral pulvinar (Pdm)], and three areas of extrastriate cortex representing early, intermediate, and late stages of cortical processing (V2, V4/PM, area 7a). A simple fixation task controlled transitions among three attentive states. The animal waited for a fixation point to appear (ready state), fixated the point until it dimmed (fixation state), and then waited idly to begin the next trial (idle state). Attentional modulation was estimated by flashing an identical, irrelevant stimulus in a neuron's receptive field during each of the three states; the three responses defined a "response vector" whose deviation from the line of equal response in all three states (the main diagonal) indicated the character and magnitude of attentional modulation. Attentional modulation was present in all visual areas except the lateral geniculate, indicating that modulation was of central origin. Prevalence of modulation was modest (26%) in pulvinar, and increased from 21% in V2 to 43% in 7a. Modulation had a push-pull character (as many cells facilitated as suppressed) with respect to the fixation state in all areas except Pdm where all cells were suppressed during fixation. The absolute magnitude of attentional modulation, measured by the angle between response vector and main diagonal expressed as a percent of the maximum possible angle, differed among brain areas. Magnitude of modulation was modest in the pulvinar (19-26%), and increased from 22% in V2 to 41% in 7a. However, average trial-to-trial variability of response, measured by the coefficient of variation, also increased across brain areas so that its difference among areas accounted for more than 90% of the difference in modulation magnitude among areas. We also measured attentional modulation by the ratio of cell discharge due to attention divided by discharge variability. The resulting signal-to-noise ratio of attention was small and constant, 1.3 +/- 10%, across all areas of pulvinar and cortex. We conclude that the pulvinar, but not the lateral geniculate, is as strongly affected by attentional state as any area of visual cortex we studied and that attentional modulation amplitude is closely tied to intrinsic variability of response.