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PubMed2003

The visual pulvinar in tree shrews II. Projections of four nuclei to areas of visual cortex.

Patterns of thalamocortical connections were related to architectonically defined subdivisions of the pulvinar complex and the dorsolateral geniculate nucleus (LGN) in tree shrews (Tupaia belangeri). Tree shrews are of special interest because they are considered close relatives of primates, and they have a highly developed visual system. Several distinguishable tracers were injected within and across cortical visual areas in individual tree shrews in order to reveal retinotopic patterns and cortical targets of subdivisions of the pulvinar. The results indicate that each of the three architectonic regions of the pulvinar has a distinctive pattern of cortical connections and that one of these divisions is further divided into two regions with different patterns of connections. Two of the pulvinar nuclei have similar retinotopic patterns of projections to caudal visual cortex. The large central nucleus of the pulvinar (Pc) projects to the first and second visual areas, V1 and V2, and an adjoining temporal dorsal area (TD) in retinotopic patterns indicating that the upper visual quadrant is represented dorsal to the lower quadrant in Pc. The smaller ventral nucleus (Pv) which stains darkly for the Cat-301 antigen, projects to these same cortical areas, with a retinotopic pattern. Pv also projects to a temporal anterior area, TA. The dorsal nucleus (Pd), which densely expresses AChE, projects to posterior and ventral areas of temporal extrastriate cortex, areas TP and TPI. A posterior nucleus, Pp, projects to anterior areas TAL and TI, of the temporal lobe, as well as TPI. Injections in different cortical areas as much as 6 mm apart labeled overlapping zones in Pp and double-labeled some cells. These results indicate that the visual pulvinar of tree shrews contains at least four functionally distinct subdivisions, or nuclei. In addition, the cortical injections revealed that the LGN projects topographically and densely to V1 and that a significant number of LGN neurons project to V2 and TD.

Acetylcholinesterase↗
PubMed2004

Retinal projections to the lateral posterior-pulvinar complex in intact and early visual cortex lesioned cats.

In intact cats, it is generally considered that the lateral posterior-pulvinar complex (LP-pulvinar) does not receive direct retinal terminals, with the exception of the retino-recipient zone known as the geniculate wing. There is, however, some evidence that early lesions of the visual cortex can occasionally induce the formation of novel retinal projections to the LP nucleus. Given the importance of knowing the connectivity pattern of the LP-pulvinar complex in intact and lesioned animals, we used the B fragment of cholera toxin, a sensitive anterograde tracer, to reinvestigate the retinal projections to the LP-pulvinar in normal cats and in cats with early unilateral lesions of the visual cortex (areas 17 and 18). Immunohistochemical localization of the toxin was performed to show the distribution and morphology of retinofugal terminals. A direct bilateral but predominantly contralateral retinal projection reached the caudal portion of LPl and LPm in the form of patches located mainly along its dorsomedial surface and many scattered terminals. The distribution of retinal projections to LP-pulvinar in intact and operated cats did not differ. Contrary to what had been previously reported, we found no evidence for lesion-induced sprouting of retinal axons in these higher-order thalamic nuclei. Retinal input to the LP-pulvinar might modulate visual responses driven by primary visual cortex or superior colliculus.

Animals↗
PubMed2001

Global motion integration in the cat's lateral posterior-pulvinar complex.

Our laboratory previously showed that thalamic neurons in an extrageniculate nucleus, the lateral posterior-pulvinar complex (LP-pulvinar) could perform higher-order neuronal operations that had until then only been attributed to higher-level cortical areas. To further assess the role of the thalamus in the analysis of complex percepts, we have investigated whether neurons in the LP-pulvinar complex can signal the direction of motion of random-dot kinematograms wherein the individual elements of the pattern do not provide coherent motion cues. Our results indicate that a subset of LP-pulvinar cells can integrate the displacement of individual elements into a global motion percept and that their large receptive fields permit the integration of motion for elements separated by large spatial intervals. We also found that almost all of the global motion-sensitive neurons were not systematically pattern-motion-selective when tested with plaid patterns. The results indicate that LP-pulvinar cells can perform the higher-level spatio-temporal integration required to detect the global displacement of objects in a complex visual scene, further supporting the notion that extrageniculate thalamic cells are involved in higher-order motion processing. Furthermore, these results provide some evidence that there may be specialized mechanisms for processing different types of complex motion within the LP-pulvinar complex.

Action Potentials↗
PubMed2003

Low medial and lateral right pulvinar volumes in schizophrenia: a postmortem study.

OBJECTIVE: In this study, the volume and neuronal number of the pulvinar thalamic nucleus in schizophrenia patients were measured. METHOD: The authors examined medial and lateral pulvinar nuclei bilaterally in 27 patients with schizophrenia and 28 normal comparison subjects. RESULTS: In the comparison subjects, the medial pulvinar was larger on the right. The right but not left pulvinar nuclei were smaller in the schizophrenia patients than in the comparison subjects. The number of neurons showed similar trends. CONCLUSIONS: These findings confirm low pulvinar volume in schizophrenia and show that it affects both medial and lateral nuclei. The lateralized findings may reflect pulvinar connections with asymmetrical neocortical regions and their asymmetrical involvement in schizophrenia.

Aged↗
PubMed1985

Anatomical evidence for medial pulvinar connections with the posterior cingulate cortex, the retrosplenial area, and the posterior parahippocampal gyrus in monkeys.

Reciprocal connections between the medial pulvinar and the limbic neocortex in monkeys were demonstrated by means of tritiated amino acid injections in the medial pulvinar and the cingulate cortex, and HRP injections in the medial pulvinar. It appears that the medial nucleus of the pulvinar sends projection fibres to the posterior cingulate gyrus (area 23), the retrosplenial area, and the posterior parahippocampal gyrus (areas TH and TF). The labeled terminals were concentrated in two bands, one in the deeper part of layer III and in layer IV, and the other in layer I. These projections were observed to be reciprocal, and the cortical afferent fibers to the medial pulvinar were found to originate from the deep layers of the cortex. The medial nucleus of the pulvinar was already known to be connected with the prefrontal cortex and with the inferior parietal lobule. Since this nucleus is now demonstrated to be connected with the posterior limbic neocortex, it is envisaged as being the thalamic counterpart of a cortical triad (prefrontal, parietal, and limbic) involved in modulating directed attention.

Animals↗
PubMed1990

Saccadic eye movements following kainic acid lesions of the pulvinar in monkeys.

Behavioral and anatomical experiments have suggested that the pulvinar might play a role in the generation of saccadic eye movements to visual targets. To test this idea, we trained monkeys to make visually-guided saccades by requiring them to detect the dimming of a small target. We used three different saccade paradigms. On single-step trials, saccades were made from a central fixation point (FP) to a target at 12, 24 or 36 degrees to the left or right. On overlap trials, the FP remained lit during presentation of a target at 12 or 24 degrees. On double-step trials, the target stepped first to 24 degrees, and then back to 12 degrees on the same side. Animals were trained to criterion, received kainic acid lesions of the pulvinar, and were retested on all three tasks. The lesions were very large, destroying almost all of the visually responsive pulvinar. They also encroached on the lateral geniculate nucleus, thereby producing small foveal scotomas, and this resulted in some behavioral changes, including difficulty in maintaining fixation on the target and in detecting its dimming. Results on the saccade tests suggest that the pulvinar is not crucial for initiation of saccadic eye movements. Saccade latency and amplitude were unimpaired on both single-step and overlap trials. Saccadic performance was also normal on double-step trials. In a second experiment, we measured the average length of fixations during spontaneous viewing of a complex visual scene. Fixation lengths did not differ from those of unoperated control monkeys. We suggest that the neglect, increased saccadic latencies, and prolonged fixations attributed to pulvinar damage in previous studies were probably the result instead of inadvertent damage to tectal afferents. The present results, together with single unit data, point to a role for the pulvinar not in the generation of saccades, but rather in the integration of saccadic eye movements with visual processing.

Animals↗
PubMed1983

Electrophysiological and anatomical demonstration of an overlapping striate and tectal projection to the lateral posterior-pulvinar complex of the cat.

Extracellular unit activity was recorded in the lateral posterior (LP)-pulvinar complex. The responses of 254 neurons after electrical stimulation of the central-paracentral part of cortical area 17 and of 84 neurons after stimulation of both area 17 and the superior colliculus (SC) were investigated. Neurons in the LP-pulvinar complex responded to area 17-stimulation with excitatory-inhibitory effects; in some cases only with inhibition. Neurons affected by striate stimulation were found in the caudal region of the complex in a region that extended widely into the medial part of the lateralis posterior nucleus (LPm), the so-called tectorecipient part of the lateral posterior nucleus. Accordingly, 26 of the 84 neurons in which electrical stimulation of area 17 and of the SC was tested, were found to react to both types of stimulation. Cells responding only to SC-stimulation were found in the ventral region of the anterior LP-Pulvinar complex. Anatomical studies supported the finding that striate and tectal inputs overlap considerably in the LP-pulvinar complex. After depositing horseradish peroxidase (HRP) into various regions of the LP-pulvinar complex, retrogradely labeled cells were found in area 17 (layer V) as well as in the superficial layers of the SC. These results were confirmed by orthograde transport autoradiography after injection of labeled amino-acids into area 17. Our findings indicate that cortical and collicular inputs into the caudal part of the LP-pulvinar complex overlap considerably and that, in these overlapping regions, individual neurons may receive converging afferent excitation from both regions.

Animals↗
PubMed1977

The morphology and laminar distribution of cortico-pulvinar neurons in the rhesus monkey.

Using autoradiography and the horseradish peroxidase method, the morphology and laminar distribution of cortico-pulvinar neurons and the reciprocity of connections between pulvinar and cortex were examined in five Rhesus monkeys which had received medial, lateral and inferior pulvinar nucleus injections of both tritiated amino acids and horseradish peroxidase. Cortico-pulvinar neurons were identified in one heterotypical cortical area (area 17) and in many homotypical areas in frontal (areas 45, 46, 11, 12), parietal (5, 7), occipital (18, 19) and temporal (20, 21, 22) lobes. The cortico-pulvinar neurons were pyramidal in shape and ranged in size from small to large. In heterotypical cortex they were found in layers V and VI whereas in area 17 they were found only in layer Vb. Reciprocal connections between pulvinar and cortex were a feature of homotypical but not heterotypical cortex.

Animals↗
PubMed1979

Topographical origin and ganglion cell type of the retino-pulvinar projection in the cat.

In order to examine the pattern of the retino-pulvinar projection in the cat, the existence of which has been recently demonstrated using autoradiographic fiber tracing technique, a small amount of horseradish peroxidase (HRP) was injected into the lateral part of the pulvinar nucleus at various rostocaudal levels. The retrogradely labeled ganglion cells were analyzed in terms of their topographical location and cell size, as seen inretinal whole mounts. The results were compared with those obtained following injections into the lateral geniculate nucleus. Retrogradely labeled cells were found in the retina bilaterally after injections of HRP into the pulvinar nucleus. Pulvinar injections produced labeling of retinal cells in the nasal half of the retina contralaterally, and in the temporal half ipsilaterally. The labeled cells were diffusely distributed in a retinotopically organized fashion. The representation of the area centralis in the retino-pulvinar projection is displaced rostrally as compared with the retino-geniculate projection. All labeled cells after pulvinar injections were medium to small size and no large cells were encountered.

Animals↗
PubMed1980

Pulvinar neuron responses to spontaneous and trained eye movements and to light flashes in squirrel monkeys.

Single unit responses were recorded in the pulvinar nucleus of the squirrel monkey (Saimiri sciureus) while awake and alert, with the head fixed but free to make eye movements. Peri-saccadic time histograms revealed only post-saccadic responses to eye movements made spontaneously in the dark or in a uniformly illuminated field (Ganzfeld), or to trained eye movements made in response to a spot of light in the periphery. Of 120 pulvinar neurons that responded to eye movements in the light, approximately one-quarter was responsive to eye movements in the dark. In one group of monkeys about half of the eye movement-responsive cells in the light were responsive to light flash in the dark; in another group approximately half of the eye movement-responsive cells in the light were also responsive to trained eye movements. Of particular interest is the fact that pulvinar neurons responsive to eye movements were located within a restricted region oriented vertically and cutting across lateral and inferior pulvinar, rostrally, and lateral and medial pulvinar, caudally. The results have been interpreted as supporting the concept of convergence and integration of visual and oculomotor input in the pulvinar and also its potential role in the mediation of visual attention.

Animals↗
PubMed1988

Connections to the lateral posterior-pulvinar thalamic complex from the reticular and ventral lateral geniculate thalamic nuclei: a topographical study in the cat.

The projections from the reticular thalamic nucleus and the ventral lateral geniculate nucleus to the lateral posterior-pulvinar thalamic complex were studied in the adult cat using the retrograde transport of horseradish peroxidase. Small, stereotaxically guided injections of the enzyme were placed in the various nuclei of this complex, including the pulvinar, lateralis intermedius oralis, lateralis intermedius caudalis, lateralis posterior lateralis, lateralis posterior medialis and lateralis medialis nuclei. The distribution of labeled neurons indicates that these nuclei receive topographically organized projections from the reticular and ventral lateral geniculate nuclei. The pulvinar nucleus receives only very scarce projections from the reticular thalamic nucleus originating in its posterodorsal and posteroventral sectors. The reticular projection to the nucleus lateralis intermedius oralis is even sparser. The nuclei lateralis intermedius caudalis, lateralis posterior lateralis and lateralis posterior medialis receive substantial projections from the suprageniculate sector of the reticular thalamic nucleus. The nucleus lateralis medialis receives an abundant projection from the three sectors (suprageniculate, pregeniculate and infrageniculate) of the reticular thalamic nucleus. Except for the lateralis intermedius caudalis, all nuclei of the lateral posterior-pulvinar complex receive consistent projections from the ventral lateral geniculate nucleus, the nucleus lateralis medialis receiving the densest one. Our findings suggest that visual, auditory, somatosensory, motor and limbic impulses from thalamic nuclei and from primary sensory and association cortical areas modulate the activity of the nucleus lateralis medialis via the reticular thalamic nucleus. The remaining nuclei of the lateral posterior-pulvinar complex are mainly modulated by sectors of the reticular thalamic nucleus that receive afferent connections from visual structures. The intrathalamic projections arising from the ventral lateral geniculate nucleus may be the way through which visuomotor inputs reach the different components of the lateral posterior-pulvinar thalamic complex.

Animals↗
PubMed2000

The pulvinar sign on magnetic resonance imaging in variant Creutzfeldt-Jakob disease.

BACKGROUND: There is a need for an accurate non-invasive diagnostic test for variant Creutzfeldt-Jakob disease (vCJD). We investigated the sensitivity and specificity of bilateral pulvinar high signal on magnetic resonance imaging (MRI) for the diagnosis of vCJD. METHODS: MRI from patients with vCJD and controls (patients with suspected CJD) were analysed. Scans were reviewed on two separate occasions by two neuroradiologists and scored for the distribution of changes, and likely final diagnosis. Scans from vCJD cases were reassessed to reach a consensus on all abnormalities. FINDINGS: We analysed 36 patients and 57 controls. vCJD patients were correctly identified based on bilateral pulvinar high signal in 29 of 36 and 32 of 36 cases on the first assessment by the two radiologists, and 32 of 36 and 31 of 36 on their second assessment. Bilateral increased pulvinar signal was identified in one of 57 and one of 57 controls on the first assessment and two of 57 and three of 57 controls on the second assessment. These reported changes in controls were graded as minimal/equivocal in six of seven patients and moderate in one (<0.5% of all control assessments). 80% of the assessments in vCJD cases were graded as moderate or substantial. On consensus review, 28 of 36 cases and none of 57 controls had prominent bilateral pulvinar signal-sensitivity 78% (95% CI 60-90%) and specificity 100% (95% CI 94-100%). Other common MRI features of vCJD were medial thalamic and periaqueductal grey matter high signal, and the notable absence of cerebral atrophy. Pulvinar high signal correlated with histological gliosis. INTERPRETATION: In the appropriate clinical context the MRI identification of bilaterally increased pulvinar signal is a useful non-invasive test for the diagnosis of vCJD.

Adolescent↗
PubMed1994

Lesions in the pulvinar in patients with Alzheimer's disease.

The pulvinar nucleus of the thalamus has been implicated in visual attention and in the control of eye movements, and may also help mediate attention-dependent tasks in the auditory and somatosensory systems. Consequently, a hypothetical disruption of functions mediated by the pulvinar may help understand common visual and nonvisual symptoms in patients with Alzheimer's disease. To test this hypothesis, the pulvinar nuclei of nine patients with histologically confirmed Alzheimer's disease and twelve young (9-28 years of age) and age-matched controls (without dementia and with non-Alzheimer dementias) were examined using a battery of histopathological methods. All patients with Alzheimer's disease had numerous amyloid plaques and some neuritic plaques throughout the various subdivisions of the pulvinar. Neuropil threads were also present among the senile plaques, and neurofibrillary tangles occurred in clusters throughout the nucleus. Control patients with and without dementia had very few Alzheimer lesions, which should be expected in normal elderly individuals. The lesions in the pulvinar may help understand some of the clinical manifestations of Alzheimer's disease, including deficits in attention and in a host of visual disturbances. Such deficits may be the result of involvement of reciprocal thalamo-cortical and corticothalamo-cortical circuits that appear to be critical for the function of a vast expanse of association cortices interconnected with the pulvinar, which are heavily laden with lesions themselves.

Adolescent↗
PubMed1999

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

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

Age Factors↗
PubMed2000

Visual cortical projections and chemoarchitecture of macaque monkey pulvinar.

We investigated the patterns of projections from the pulvinar to visual areas V1, V2, V4, and MT, and their relationships to pulvinar subdivisions based on patterns of calbindin (CB) immunostaining and estimates of visual field maps (P(1), P(2) and P(3)). Multiple retrograde tracers were placed into V1, V2, V4, and/or MT in 11 adult macaque monkeys. The inferior pulvinar (PI) was subdivided into medial (PI(M)), posterior (PI(P)), central medial (PI(CM)), and central lateral (PI(CL)) regions, confirming earlier CB studies. The P(1) map includes PI(CL) and the ventromedial portion of the lateral pulvinar (PL), P(2) is found in ventrolateral PL, and P(3) includes PI(P), PI(M), and PI(CM). Projections to areas V1 and V2 were found to be overlapping in P(1) and P(2), but those from P(2) to V2 were denser than those to V1. V2 also received light projections from PI(CM) and, less reliably, from PI(M). Neurons projecting to V4 and MT were more abundant than those projecting to V1 and V2. Those projecting to V4 were observed in P(1), densely in P(2), and also in PI(CM) and PI(P) of P(3). Those projecting to MT were found in P(1)- P(3), with the heaviest projection from P(3). Projections from P(3) to MT and V4 were mainly interdigitated, with the densest to MT arising from PI(M) and the densest to V4 arising from PI(P) and PI(CM). Because the calbindin-rich and -poor regions of P(3) corresponded to differential patterns of cortical connectivity, the results suggest that CB may further delineate functional subdivisions in the pulvinar.

Animals↗
PubMed2006

Long-range interneurons within the medial pulvinar nucleus of macaque monkeys.

Like other thalamic nuclei, the primate pulvinar is considered not to have long-range intrinsic connections, either excitatory or inhibitory. Injections of biotinylated dextran amine (BDA) in the medial pulvinar, however, reveal retrogradely filled neurons up to 2.0 mm from the injection edge. Serial section reconstruction (n = 18) confirmed that retrogradely filled neurons projected to the injection site and showed that they had additional long-range collaterals within the posterior pulvinar. Arrays of small, beaded terminations occurred in multiple foci along the collaterals. Terminal arrays were up to 1.0 mm in length; foci were separated by about 0.7 mm. Somata were large (average area = 220 microm2), and dendritic arbors were radiate and also large (about 1.0 mm in diameter), but without either the appendages of classical interneurons or the hairlike spines characteristic of radiate pulvinocortical projection neurons. Double labeling for BDA and parvalbumin (PV) or BDA and gamma-aminobutyric acid (GABA) indicated that these large neurons were positive for both PV and GABA. Double labeling for PV and GABA, or PV and glutamic acid decarboxylase 67 (GAD67) revealed a small number of similarly large neurons in the posterior pulvinar that were positive for both substances. Thus, we propose that these neurons are a novel class of inhibitory interneuron, longer range than the classic thalamic local circuit interneurons. Future questions include how these neurons relate to other inhibitory systems and specific postsynaptic populations and whether they are located preferentially within the posterior pulvinar, possibly related to the multimodal character of this thalamic region.

Animals↗
PubMed2006

5HTTLPR polymorphism and enlargement of the pulvinar: unlocking the backdoor to the limbic system.

BACKGROUND: The 5HTTLPR genetic variant of the serotonin transporter (SERT), which consists of a long (SERT-l) and short (SERT-s) allele, has emerged as a major factor influencing emotional behavior and brain anatomy. The pulvinar nucleus of the thalamus projects to important limbic nuclei including the amygdala and cingulate cortex, is involved in the processing of stimuli with emotional content, and contains an abundance of SERT. METHODS: Stereological methods were used to measure pulvinar neuron number in postmortem tissue from major depressive disorder (n = 11), bipolar disorder (n = 11), schizophrenia (n = 12), and control (n = 15) specimens from the Stanley Foundation Neuropathology Consortium. The effect of SERT genotype on pulvinar volume and neuron number was investigated by using analysis of covariance. RESULTS: Analysis of covariance with diagnosis, SERT genotype, age, hemisphere, postmortem interval, and time-in-formalin covariates identified a 20% increase in pulvinar neuron number and volume in SERT-ss subjects. CONCLUSIONS: The elevated number of pulvinar neurons in subjects with a SERT-ss genotype may serve to enhance subcortical input of emotionally relevant stimuli to the limbic system, providing a mechanism for the 5HTTLPR genetic variant to affect predisposition to conditions such as major depression.

Alleles↗
PubMed2005

Response to visual threat following damage to the pulvinar.

We present a unique case demonstrating contributions of the pulvinar in response to visual threat. Substantial evidence demonstrates that the amygdala contributes to the emotion of fear and the response to threat. Traditionally, two routes to amygdala activation have been distinguished: a "slow cortical" route through visual and association cortex and a "fast subcortical" route through the thalamus. The pulvinar nucleus of the thalamus is well connected to the amygdala, suggesting that pulvinar damage might interfere with amygdala activation and response to threat. We tested this possibility in patient SM, who suffered complete loss of the left pulvinar. We measured interference from threatening images on goal-directed behavior. In SM's ipsilesional field, threatening images slowed responses more than pleasant images did. This interference decreased rapidly over time. In contrast, in SM's contralesional field, interference from threatening images was initially absent and then increased rather than decreased over time. Processing through the pulvinar therefore plays a significant role in generating response to visual threat. We suggest that, with disruption of the subcortical route to the amygdala, briefly presented images were not fully processed for threat. The reemergence of interference over time may reflect contributions of a slower route.

Amygdala↗