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PubMedDate not supplied

Connectional and neurochemical subdivisions of the pulvinar in Cebus monkeys.

Based on cytoarchitectonic criteria, the primate pulvinar nucleus has been subdivided into medial (PM), lateral (PL), and inferior (PI) regions. However, these subdivisions show no correlation with those established by electrophysiological, immunocytochemical, or neuroanatomical tracer studies. In this work, we studied the connections of the pulvinar nucleus of Cebus monkey with visual areas V1, V2, V4, MT, and PO by means of retrograde fluorescent tracers injected into these areas. Based on the projection zones to cortical visual areas, the visual portion of the pulvinar of Cebus monkey was subdivided into three subregions: P1, P2, and P3, similar to those described in the macaque (Ungerleider et al., 1984). In Cebus, P1 includes the centrolateral portion of traditionally defined PI and adjacent portion of PL. P2 is located in the dorsal portion of PL and P3 includes the medial portion of PI and extends dorsally into adjacent PL and PM. In addition, we studied the histology of the pulvinar using multiple criteria, such as cytoarchitecture and myeloarchitecture; histochemistry for cytochrome oxidase, NADPH-diaphorase, and acetylcholinesterase; and immunocytochemistry for two calcium-binding proteins, calbindin and parvalbumin, and for a neurofilament recognized by the SMI-32 antibody. Some of these stains, mainly calbindin, showed additional subdivisions of the Cebus pulvinar, beyond the traditional PI, PL, and PM. Based on this immunohistochemical staining, the border of PI is moved dorsally above the brachium of the superior colliculus and PI can be subdivided in five regions (PI(P), PI(M), PI(C), PI(L), and PI(LS)). Regions P1, P2, and P3 defined based on efferent connections with cortical visual areas are not architectonically/neurochemically homogeneous. Rather they appear to consist of further chemoarchitectonic subdivisions. These distinct histochemical regions might be related to different functional modules of visual processing within one connectional area.

Acetylcholinesterase↗
PubMed1979

The inferior pulvinar complex in owl monkeys: architectonic subdivisions and patterns of input from the superior colliculus and subdivisions of visual cortex.

Patterns of connections with other visual structures and architectonic characteristics were used to subdivide the inferior pulvinar complex of owl monkeys into three distinct nuclei termed the central inferior pulvinar, IPc, the medial inferior pulvinar, IPm, and the posterior inferior pulvinar, IPp. IPc occupies about 70%; IPm about 20%, and IPp about 10% of the inferior pulvinar complex. Encapsulating fiber bands distinguish the boundaries of the three nuclei. IPm is also identified by a much greater packing density of neurons than IPc and IPp. Both IPp and IPc receive input from the superior colliculus, but the terminations in IPp are denser. Visual cortical Areas 17, 18, MT, DM, M and PP (Allman and Kaas, '76) project to IPc and IPm in patterns that indicate that central vision is represented dorsorostrally and peripheral vision ventrocaudally in both nuclei. Terminations in IPm from Area MT are particularly dense. None of these visual areas projects to IPp. Rather, input to IPp appears to originate in cortex rostral to Area MT in the temporal lobe.

Animals↗
PubMed1979

The structural organization of the inferior and lateral subdivisions of the Macaca monkey pulvinar.

Previous light microscopic studies of Macaca pulvinar have demonstrated that both the inferior and adjacent portion of the lateral pulvinar subdivisions are reciprocally connected to the entire occipital lobe, including striate cortex. They differ in that inferior but not lateral pulvinar receives a projection from the superficial layers of the superior colliculus. In this study, the internal organization of these two subdivisions in compared by relating light microscopic Golgi morphology to the synaptic organization observed by electron microscopy. The Golgi impregnated neurons in inferior and lateral pulvinar are typical of other thalamic nuclei and are not qualitatively different in the two subdivisions. Projections neurons (PN) vary in cell body (15--40 micrometers) and dendritic tree (150--600 micrometers) diameters but bear the same varieties of dendritic appendages; spine-like, hair-like, and knot-like. Local circuit neurons (LCN) have smaller cell body diameters (10--20 micrometers) but can have very large dendritic field diameters (150--600 micrometers). They are best distinguished from PNs by their elaborate dendritic appendages, which have been identified as pre-synaptic dendrites in the EM. LCN axons are infrequently seen. In the EM both subdivisions contain four types of synaptic terminals. RS and RL terminal both contain round symaptic vesicles and make asymmetric synaptic contacts, but are subdivided on the basis of small (RS = 0.09 micrometers) versus large (RL = 2.2 micrometers) cross sectional diameters and organelle content. RLs contact larger caliber dendrites and frequently form synaptic complexes with presynaptic dendrites of LCNs, while RSs contact fine caliber dendrites and rarely take part in synaptic complexes. F terminal and P boutons both contain flat and pleomorphis vesicles and make symmetric synaptic contacts. They are characterized by vesicle number and cytoplasmic density. Fs are infrequently observed in pulvinar compared to P boutons and are of uncertain origin. P boutons can be equated with LCN dendritic appendages and have been identified as pre-synaptic dendrites. The quantitative distribution of each type is very similar in both subdivisions, avveraging RS 85%, RL 5%, F 0.3%, P 8% and unidentified 2%.

Animals↗
PubMed1980

Cortical connections of the pulvinar nucleus in Galago.

In the first series of experiments, small amounts of HRP were injected into areas 17, 18, and 19 and each of the cytoarchitectonic areas of the temporal lobe. The resulting distributions of labeled cells fell into a number of distinctive classes. For example, after injecting the temporal anterior area (Ta), the labeled cells occupied a band on the ventral border of the inferior division of the pulvinar complex; after injecting the temporal ventral area (Tv), the labeled cells were concentrated in the medial extremity of the superior division. In spite of the distinctiveness of these different distributions, there was evidence that in areas that we know to contain a representation of the visual field, the injection of the same part of the field led to labeled cells in the same part of the pulvinar nucleus. Thus, when the representation of the center of the field was injected in areas 17 or 18 or Tm (the temporal middle area), labeled cells were found in the dorsal part of the rostral half of the inferior division. The distinctiveness of the different distributions did not obscure certain features common to all experiments: labeled cells were always found in both subdivisions of the pulvinar complex, and there was always continuity between the population of labeled cells in the inferior division and the population of cells in the superior division. Wherever the site of the injection in the extrastriate region, some labeled cells were found in the causal half of the inferior division. Since the caudal half of the inferior division corresponds approximately to the tecto-recipient zone as defined earlier, the entire temporal lobe, except for the auditory areas, is visual cortex. Evidence was also found for an overlap in the striate cortex between the projections of the lateral geniculate body and the pulvinar nucleus. In conclusion, the pulvinar complex projects to a vast area of cortex, including all of the occipital lobe and most of the lateral surface of the temporal lobe. This entire region, which comprises a good part of the whole of the neocortex, can be regarded as visual cortex. A second series of experiments involved cortical injections of tritiated amino acids. The results showed that the projections from the thalamus to cortex were precisely reciprocated by descending projections from cortex to thalamus. The results also served as a way of confirming the results of the first set of experiments.

Animals↗
PubMed1983

Sensorimotor unit activity related to intention in the pulvinar of behaving Cebus Apella monkeys.

Previous observations made in our laboratory in a waking behaving Cebus Apella monkey revealed that neurons of the Pulvinar nucleus of the thalamus discharged preferentially in relation to intentional movements of the limbs and eyes. We give here a description of further observations made in two waking, behaving Cebus Apella monkeys trained to perform two tasks: in the first to make saccadic eye movements to eccentrically placed visual targets; in the second, to make projection movements of the arm and hand to touch targets within arm's reach. The electrical activity of thalamic neurons was recorded extracellularly and records were made simultaneously of the horizontal eye movements and of tasks events. Four-hundred-sixty-five neurons were studied: of these, the activity of 272 could be correlated with behavioral events, while the remaining 193 could not be correlated or classified in this manner. The cells identified were classed in five groups, as follows: (1) neurons active during attentive fixation of a target, but which did not respond to our ordinary visual test stimuli; (2) neurons active during projection movements of the arm or manipulation with the hand, but which were not active during casual movements of the hand or arm, and which were not activated by passive somatic sensory stimuli; (3) those active before, during or after evoked saccadic movements of the eyes, but which were not activated by our testing visual stimuli; (4) neurons active during tracking movements of the eyes, or during projection movements of the arm, alone, but which discharged maximally when these two events occurred simultaneously; and, (5) neurons active during both saccadic movements of the eyes and during projection movements of the arm. We regularly observed, for each of these classes of neurons of the Pulvinar, that optimal correlated activity depended upon the intentional nature of the associated behavioral events, and the animal's attention to them. We conclude that there exists at the level of the Pulvinar a neural correlate of certain evolving behavioral events, and particularly of intentional activity such as the projection of the arm or the direction of gaze towards targets of interest in the immediately surrounding visual environment. The regions of the Pulvinar containing neurons with these properties are reciprocally related to association areas of the Neocortex known to contain neurons with similar properties. It can then be concluded from both anatomical and electrophysiological observations that the Pulvinar is an essential part of the system controlling these complex behavioral events.

Afferent Pathways↗
PubMed1981

Differential responsiveness of cells in the visual zones of the cat's LP-pulvinar complex to visual stimuli.

Multiple visual field representations are contained within the feline LP-pulvinar complex; regions differentiated by their afferent and efferent connectivity patterns as the striate-, tecto- and retino-recipient zones. Cell responses from these visuotopic zones were investigated in immobilized cats under N2O/O2 supplemented with pentobarbitone or Althesin, using spot, bar and textured stimuli. Response fields recorded within the LP-pulvinar complex were classified as diffuse, concentric, movement-, direction- or orientation-sensitive. Concentric receptive fields were further classified as sustained (X), transient (Y) or tonic/phasic W-cells. Direction- and movement-sensitive cells predominated in the striate- and tecto-recipient zones, respectively. Motion of noise fields, or noise bars against an identical stationary noise background elicited vigorous responses from cells in the striate zone, many showing a preference for noise stimuli. In contrast, cells from the tectal zone and other divisions of the LP-pulvinar complex were insensitive to noise. The retino-recipient zone at the lateral margin of the pulvinar nucleus was characterized by cells with concentric receptive fields, the majority exhibiting properties similar to W-cells in the LGNd. The evidence supports the notion of functional subdivision within the LP-pulvinar complex corresponding to the visuotopically organized regions defined by their connectivity patterns. Consideration of the retino-recipient zone as an extension of the LGNd-MIN complex is discussed.

Animals↗
PubMed1984

Effects of kainic acid and radiofrequency lesions of the pulvinar on visual discrimination in the monkey.

Monkeys with thermocoagulation or kainic acid lesions of the pulvinar and unoperated control monkeys were tested in two tasks: pattern discrimination retention and color discrimination learning in which the stimuli were located at the response sites or were separated spatially from them (S-R separation). The monkeys with kainic acid pulvinar lesions were mildly impaired in retention of the pattern discrimination, but were unimpaired in the color discrimination tasks with or without the S-R separation. The monkeys with thermocoagulation lesions, like monkeys with superior colliculus lesions in a prior study, were severely impaired in performing one of the color discrimination tasks with S-R separation. These findings suggest that: (a) the inferior pulvinar, unlike the superior colliculus, does not contribute to the performance of discriminations involving S-R separation; and (b) corticotectal projections traversing the pulvinar and destroyed by the thermocoagulation lesions are crucial to the performance of discriminations involving S-R separation. The results of an earlier experiment also suggested that interruption of corticotectal fibers passing through the pulvinar impairs performance in another task sensitive to superior colliculus lesions--spatial localization of light flashes. Thus, corticotectal projections may be crucial for the contribution of the colliculus to performance in a variety of visual tasks.

Animals↗
PubMed1989

Visual receptive fields in the striate-recipient zone of the lateral posterior-pulvinar complex.

The lateral posterior (LP)-pulvinar complex of the cat is known to contain multiple visual areas. In the present study, we examined the receptive field properties of single neurons isolated in the lateral division of this complex (the LPI). The LPI is designated the striate-recipient zone because it is the only region of the LP-pulvinar receiving cortical projections from areas 17 and 18. The recordings revealed that the striate-recipient zone of LP comprises 2 subareas, which we have termed LPI-1 and LPI-2. In the main segment (LPI-1), virtually all cells responded securely to visual stimuli. The vast majority of these neurons were binocular, with relatively small and well-defined receptive fields. More than half of the cells were found to be directionally selective, and almost this many were orientation specific. The orientation tuning of these cells was found to be quite precise, comparable to complex cells in area 17. In contrast, in the small dorsolateral segment of the striate-recipient zone (the LPI-2), a substantial proportion of cells could not be visually activated. Here, the visual cells had very large receptive fields, and relatively few were direction or orientation selective. The LPI-2 receives subcortical inputs from the superficial layers of the superior colliculus, the hypothalamus, and cerebellum, while the LPI-1 is innervated only by cortical axons. It is suggested that the subcortical connections of the LPI-2 account for the differences in the response properties of the 2 striate-recipient areas. The present results, in conjunction with our previous findings on the principal tectorecipient zone (Chalupa et al., 1983), permit 2 generalizations regarding the functional organization of the cat's LP-pulvinar complex. First, there are clear differences among the visual areas of the LP-pulvinar in the cellular processing of visual information. Second, these functional differences can be related to the principal sources of visual input to the various divisions of the LP-pulvinar.

Afferent Pathways↗
PubMedDate not supplied

[Interaction of recovery cycles of evoked potentials in the cerebral cortex of the cat in response to stimulation of the superior colliculi and pulvinar].

Similar character of recovery cycles of evoked potentials in the visual cortex to electric stimulation of the superior colliculi (SC) and pulvinar was found in chronic experiments on alert cats irrespective of stimuli presentation order. In the association cortex preceding SC stimulation facilitated the response to test stimulation of pulvinar almost at all delays between the stimuli. If the pulvinar stimulation was applied as a conditioned stimulus, then the response to SC stimulation under intervals of 20-200 ms was depressed. The obtained data point to equivalence of the inputs from SC and pulvinar to the visual cortex, to different informational value of inputs from SC to the association and visual cortex, and to mutual function dependence of the inputs from SC and pulvinar to the association cortex.

Animals↗
PubMed2005

Neurochemical organization of chimpanzee inferior pulvinar complex.

The pulvinar of primates, which connects with all visual areas, has been implicated in visual attention and in control of eye movements. Recently, five separate neurochemical subdivisions of a region termed the inferior pulvinar complex have been identified in monkeys (Gray et al. [1999] J Comp Neurol 409:452-468; Gutierrez et al. [1995] J Comp Neurol 363:545-562), and comparable subdivisions have been mapped in humans (Cola et al. [1999] NeuroReport 10:3733-3738). In the present study, we investigated the inferior pulvinar of the chimpanzee (Pan troglodytes), the closest evolutionary relative of humans, using cytochrome oxidase (CO) and acetylcholinesterase (AChE) histochemistry, and immunocytochemistry for calbindin. Each staining method demarcated five histochemical zones corresponding, from medial to lateral, to the posterior (PI(P)), medial (PI(M)), central PI(C)), lateral (PI(L)), and the lateral-shell (PI(L-S)) divisions in monkeys. The PI(P) division stained darkly for calbindin and lightly for CO and AChE. The PI(M) division was characterized by less neuropil staining for calbindin, and by distinct, intensely stained patches of CO and AChE. PI(C) appeared lighter than adjacent divisions with CO and AChE histochemistry and was moderately stained with calbindin. PI(L) was moderately to darkly stained with each method and was adjoined by a lighter staining shell, PI(L-S). Thus, in the aspects of organization we examined, the inferior pulvinar of chimpanzees closely resembles that of humans and monkeys. This investigation provides a foundation for more detailed studies of the thalamic relationships of extrastriate cortex in apes and humans.

Acetylcholinesterase↗
PubMed2006

Overlapping visual response latency distributions in visual cortices and LP-pulvinar complex of the cat.

The visual system of the cat is considered to be organized in both a serial and parallel manner. Studies of visual onset latencies generally suggest that parallel processing occurs throughout the dorsal stream. These studies are at odds with the proposed hierarchies of visual areas based on termination patterns of cortico-cortical projections. In previous studies, a variety of stimuli have been used to compute latencies, and this is problematic as latencies are known to depend on stimulus parameters. This could explain the discrepancy between latency and neuroanatomical based studies. Therefore, the first aim of the present study was to determine whether latencies increased along the hierarchy of visual areas when the same stimuli are used. In addition, the effect of stimulus complexity was assessed. Visual onset latencies were calculated for area 17, PMLS, AMLS, and AEV neurons. Latencies were also computed from neurons in the lateral posterior (LP)-pulvinar complex given the importance of this extrageniculate complex in cortical intercommunication. Latency distributions from all regions overlapped substantially, and no significant difference was present, regardless of the type of stimulus used. The onset latencies in the LP-pulvinar complex were comparable to those seen in cortical areas. The data suggest that the initial processing of information in the visual system is parallel, despite the presence of a neuroanatomical hierarchy. Simultaneous response onsets among cortical areas and the LP-pulvinar suggest that the latter is more than a simple relay station for information headed to cortex. The data are consistent with proposals of the LP-pulvinar as a center for the integration and distribution of information from/to multiple cortical areas.

Animals↗
PubMed2006

Thalamocortical circuits: fMRI assessment of the pulvinar and medial dorsal nucleus in normal volunteers.

This fMRI study investigates the activation of the thalamic nuclei in a spatial focusing-of-attention task previously shown to activate the pulvinar with FDG-PET and assesses the connectivity of the thalamic nuclei with cortical areas. Normal right-handed subjects (eight men, eight women, average age=32 years) viewed four types of stimuli positioned to the right or left of the central fixation point (left hemifield-large letter, left hemifield-small letter display with flanking letters; right hemifield-large letter, right hemifield-small letter display with flankers). BOLD responses to small letters surrounded by flankers were compared with responses to large isolated letters. To examine maximum functional regional connectivity, we modeled "subject" as a random effect and attained fixed effect parameter estimates and t-statistics for functional connectivity between each of the thalamic nuclei (pulvinar, medial dorsal, and anterior) as the seed region and each non-seed voxel. Greater BOLD activation for letters surrounded by flankers than for large letters was observed in the pulvinar as anticipated and was also marked in the medial dorsal nucleus (MDN), anterior and superior cingulate (BA24 and BA24'), dorsolateral prefrontal cortex, and frontal operculum and insula. For the MDN, maximal functional connectivity was with the dorsolateral prefrontal cortex; correlations with left superior temporal, parietal, posterior frontal, and occipital regions were also observed. For the pulvinar, maximal functional connectivity was with parietal BA39; for anterior thalamus, with anterior cingulate.

Adult↗
PubMed2006

Reduction of right medial pulvinar volume and neuron number in schizophrenia.

Several studies have described a reduction of pulvinar volume and/or neuronal number in schizophrenia (SZ). In order to better localize these changes, we assessed volume and neuronal number of the pulvinar and several of its subdivisions in postmortem material from subjects with chronic SZ and subjects with no psychiatric history. Total pulvinar volume and neuronal number were significantly lower in SZ and these differences were significant in only in its medial division. The medial pulvinar interconnects various heteromodal cortical regions suggesting that these deficits may contribute to the abnormalities of higher order integrative functions characteristic of schizophrenia.

Aged↗
PubMed2001

The retinal input to calbindin-D28k-defined subdivisions in macaque inferior pulvinar.

Several studies have provided evidence for direct retinal input to the pulvinar of macaques monkeys, but there is no general agreement regarding the extent of this projection. Moreover, it is not known how retinal input correlates with chemoarchitectonic subdivisions recently recognized within the large, classical divisions of the pulvinar. The potential implications of this correlation have become more evident after reports that chemoarchitectonic subdivisions of the inferior pulvinar (PI) have specific patterns of connections with cortical visual areas. We have therefore re-examined the retino-PI projection using intraocular injections of horseradish peroxides, and correlated it with pulvinar subdivisions revealed using an antibody for calbindin-D28k. Retinal projections were found preferentially within the medial subdivision of the PI, with some involvement of the posterior and central calbindin-D28k defined subdivisions.

Animals↗
PubMed2002

Attending to the thalamus: inhibition of return and nasal-temporal asymmetry in the pulvinar.

Inhibition of return (IOR) is a mechanism whereby the attentional system favors novel locations by inhibiting already scanned ones. An important question is what the neural structures are involved. Recently, we studied a patient with damage to the superior colliculus (SC) and concluded that the SC generates IOR. However, it is possible that IOR is generated beyond the colliculus, for example, by the pulvinar. In this paper we tested three patients with unilateral damage to the pulvinar and demonstrated that the pulvinar is not necessary for IOR generation, providing additional support to the suggestion that the SC generates IOR. In addition, since we used monocular presentation, we were able to furnish behavioral evidence for nasal-temporal asymmetrical representation of visual input in the pulvinar.

Analysis of Variance↗
PubMed2003

T1 hyperintensity in the pulvinar: key imaging feature for diagnosis of Fabry disease.

BACKGROUND AND PURPOSE: Fabry disease (FD) is an inborn error of glycosphingolipid metabolism. To date, no specific neuroimaging features have been elucidated to help in making the diagnosis of this disorder. The purpose of this study was to determine whether the finding of T1 shortening in the lateral pulvinar is a useful finding in the imaging diagnosis of FD and to deduce the relationship of this finding to the pathophysiology of the disease. METHODS: We studied T1- and T2-weighted images obtained in ten patients (nine male and one female) with FD with an age range of 19-59 years. The images were examined for anatomic aberrations and areas of abnormal signal intensity (SI) in both gray matter and white matter. The SI of deep gray matter was evaluated qualitatively and semiquantitatively, relative to the SI of CSF or the genu of the corpus callosum. Gradient echo MR images and axial noncontrast CT images were available for one patient. RESULTS: Seven of 10 patients showed small areas of T2 prolongation in the white matter of the cerebral hemispheres. Despite the known propensity for vascular disease in these patients, only one had cortical infarction. Bilateral T1 shortening in the lateral pulvinar was recognized in at least seven patients, all over the age of 30 years, who also had small areas of T2 prolongation in the white matter. CT and gradient echo images in one patient revealed no evidence of calcification or metallic deposits in the pulvinar. CONCLUSION: Bilateral T1 shortening in the lateral pulvinar is a common finding in FD and may be useful in suggesting this diagnosis.

Adult↗
PubMed1981

Short- and long-term effects of neonatal and adult visual cortex lesions on the retinal projection to the pulvinar in cats.

An increased retinal projection to th pulvinar occurred in cats following neonatal ablation of visual cortical areas 17, 18, and 19. After unilateral lesions, the retinopulvinar projection visualized with autoradiography was larger and denser on the lesioned side than on the intact side. This increased projection was first recognized when the lesioned kittens were 1 week old, when increased labeling in the pulvinar was first detected. The retinopulvinar projection on the intact side of the brain decreased in size with age over the first 3 months of life. This decrease in size was not observed on the lesioned side of the brain. The absence of this decrease on the lesioned side may represent a failure of retraction of exuberant connections, but because of the active invasion by retinal fibers immediately after the lesion, observations of the normal retraction may simply be obscured. Severe cellular degeneration in the dorsal lateral geniculate occurred a few days before the changes in the retinopulvinar projection began; this loss of a postsynaptic target for many retinal axons may trigger the reorganization of input to the pulvinar retino-recipient zone (pulivnar-RRZ). After adult visual cortex lesions, no reorganization of the retinopulvinar projection took place. The modification of the projection from the retina to the pulvinar that occurs after ablation of visual cortex in the newborn cat may contribute to the sparing of visual abilities seen after these lesions.

Aging↗
PubMed1981

Projections from the superior colliculus and the neocortex to the pulvinar nucleus in Galago.

We have studied the projections from the superior colliculus and the neocortex to the pulvinar nucleus in Galago senegalensis by using the retrograde transport of horseradish peroxidase (HRP). Injections of various parts of the pulvinar complex, both the inferior and superior divisions, both the tectorecipient zone and the nontectorecipient zone as defined by Glendenning et. al. ('75), produce labeled cells in the lower tier of stratum griseum superficiale. The distribution of labeled cells in the superior colliculus varies with the locus of the injection, indicating a retinotopic projection system from the entire superior colliculus to all sectors of the pulvinar complex. These experiments also provide an opportunity to study the distribution and laminar origin of neurons giving rise to cortical descending projections. The entire visual cortex projects onto the pulvinar complex. The cells or origin can be divided into two populations--one located in layer V and the other in layer VI. In seven of the nine cases reported, the layer V population is restricted entirely or mainly to the striate area. In the two exceptional cases, the layer V population is located in the adjacent extrastriate cortex, areas 18 and 19. The difference in the layer of origin of the cortical descending fibers reflects a difference in the layer of termination of the reciprocal ascending projection. These findings identify the entire visual field as primary visual cortex. The importance of this conclusion is underscored by the fact that the visual field comprises as much as one-half of the whole neocortex.

Animals↗