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Cortical projections to the superior colliculus in the macaque monkey: a retrograde study using horseradish peroxidase.

The topical and laminar distribution of corticotectal cells, as well as their size and morphology, were studied in the macaque monkey with the horseradish peroxidase (HRP) technique. After HRP injections restricted primarily to the superficial layers of the colliculus, labelled cells were found in visual cortex (areas 17, 18, and 19) and both in the frontal eye field (area 8) and the adjacent part of premotor cortex (area 6). The clustering of labelled cells in visual cortex indicated that each of the anatomically and functionally distinct visual areas has its own set of collicular projections. When intermediate and deeper layers of the colliculus were injected, labelled cells were found also in posterior parietal cortex (area 7) where they were concentrated mainly on the posterior bank of the intraparietal fissure, in inferotemporal cortex (areas 20 and 21), in auditory cortex (area 22), in the somatosensory representation SII (anterior bank of sylvian fissure, area 2), in upper insular cortex (area 14), in motor cortex (area 4), in premotor cortex (area 6), and in prefrontal cortex (area 9). In the motor and premotor cortex, labelled cells formed a continuous band which appeared to stretch across finger-hand-arm-shoulder-neck representation. Similarly, the cluster of labelled cells in area 2 may correspond to the finger-hand representation of SII. The cortical regions not containing labelled cells were the somatosensory representation SI (areas 3, 1 and 2) and the infraorbital cortex. Labelled cells were restricted to layer V of all cortical areas except in the primary visual cortex, where labelled cells were found in both layer V and layer VI. The size spectrum of corticotectal cells ranged from 14.8 micron (average diameter) in area 17 to 27.8 micron in area 6, comprising cells as small as 8 micron and as large as 45 micron. Labelled cells in posterior parietal (area 7), in auditory (area 22), and in motor cortex (area 4) were small and distributed over only a narrow range of sizes. Those in premotor cortex (area 6) were often large and had a wide range in size distribution. The differences in size and morphology of corticotectal neurons suggest that they do not form a uniform class of neurons.

Animals↗

Retinotopic map plasticity in adult cat visual cortex is accompanied by changes in Ca2+/calmodulin-dependent protein kinase II alpha autophosphorylation.

In adult cats, the induction of homonymous binocular central retinal lesions causes a dramatic reorganization of the topographic map in the sensory-deprived region of the primary visual cortex. To investigate the possible involvement of the alpha-subunit of the calcium/calmodulin dependent protein kinase type II (alphaCaMKII) in this form of brain plasticity, we performed in situ hybridization and Western blotting experiments to analyze mRNA, protein and autophosphorylation levels of this multifunctional kinase. No differences in the mRNA or protein levels were observed between the central, sensory-deprived and the peripheral, non-deprived regions of area 17 of retinal lesion animals or between corresponding cortical regions of normal control animals. Western blotting with an alphaCaMKII threonine-286 phosphorylation-state specific antiserum consistently showed a small, albeit not significant, increase of alphaCaMKII autophosphorylation in the central versus the peripheral region of cortical area 17, and this both in normal subjects as well as in retinal lesion animals with a 3-day post-lesion survival time. In contrast, a post-lesion survival time of 14 days resulted in a alphaCaMKII autophosphorylation level that was four times higher in visually-deprived area 17 than in the non-deprived cortical region. This increased phosphorylation state is not a direct consequence of the decrease in visual activity in these neurons, because we would have expected to see a similar change at shorter or longer post-lesion survival times or in the visually deprived visual cortex of animals in which the left optic tract and the corpus callosum were surgically cut. No such changes were observed, leading to the conclusion that the phosphorylation changes observed at 14 days are related to a delayed reorganization of the retinotopic map of the striate cortex.

Animals↗

Layer-specific differential regulation of transcription factors Zif268 and Jun-D in visual cortex V1 and V2 of macaque monkeys.

To investigate intracellular mechanisms of cortical layer-specific gene regulation, we quantitatively examined the expression of two transcription factors, Zif268 and JunD, and compared their expression levels in each layer of the primary visual cortex (VI) and visual area 2 (V2) of macaque monkeys (Macaca fuscata). The brain sections were immunohistochemically stained for determination of the percentage of Zif268- or JunD-expressing neurons in the total neuronal population. We found area- and layer-specific expression of these transcription factors; Zif268 tended to be expressed at high levels in layers on the parvocellular pathway in V1, whereas JunD did not show such an expression pattern. In V1, many Zif268-immunopositive neurons were observed in layers II/III, IVC beta and VI. The percentage of Zif268-immunopositive neurons was highest in layer IVC beta and lowest in layer IVC alpha. JunD-immunopositive neurons were fewest in layer IVC beta and most abundant in layer VI. In V2, the level of expression of Zif268 was almost the same as that of JunD in layer II/II. However, layer IV of V2 tended to contain more Zif268-immunopositive neurons than JunD-immunopositive neurons, whereas layer VI contained more JunD-immunopositive neurons than Zif268-immunopositive neurons. Although it has been reported that the same extracellular signals induce both Zif268 and JunD, the present results indicate that the expression of these transcription factors is differentially controlled in each layer of the primate visual cortical areas. Furthermore, the present results suggest that these transcription factors contribute to area- and layer-specific gene regulation by mediating transmission of extracellular signals to the nucleus via different intracellular signalling pathways.

Animals↗

Selective activation of visual cortex neurons by fixational eye movements: implications for neural coding.

During normal vision, when subjects attempt to fix their gaze on a small stimulus feature, small fixational eye movements persist. We have recorded the impulse activity of single neurons in primary visual cortex (V1) of macaque monkeys while their fixational eye movements moved the receptive-field activating region (AR) over and around a stationary stimulus. Three types of eye movement activation were found. (1) Saccade cells discharged when a fixational saccade moved the AR onto the stimulus, off the stimulus, or across the stimulus. (2) Position/drift cells discharged during the intersaccadic (drift) intervals and were not activated by saccades that swept the AR across the stimulus without remaining on it. To activate these neurons, it was essential that the AR be placed on the stimulus and many of these cells were selective for the sign of contrast. They had smaller ARs than the other cell types. (3) Mixed cells fired bursts of activity immediately following saccades and continued to fire at a lower rate during intersaccadic intervals. The tendency of each neuron to fire transient bursts or sustained trains of impulses following saccades was strongly correlated with the transiency of its response to stationary flashed stimuli. For one monkey, an extraretinal influence accompanying fixational saccades was identified. During natural viewing, the different eye movement classes probably make different contributions to visual processing. Position/drift neurons are well suited for coding spatial details of the visual scene because of their small AR size and their selectivity for sign of contrast and retinal position. However, saccade neurons transmit information that is ambiguous with respect to the spatial details of the retinal image because they are activated whether the AR lands on a stimulus contour, or the AR leaves or crosses the contour and lands in another location. Saccade neurons may be involved in constructing a stable world in spite of incessant retinal image motion, as well as in suppressing potentially confusing input associated with saccades.

Action Potentials↗

[The sensitivity of the cat visual cortex neurons to cruciform figures during stimulation of the central and peripheral receptive fields].

The role of the central and peripheral zones of the receptive field (RF) in sensitivity to cruciform figures was studied in 29 neurons of the cat primary visual cortex. Separated or combined stimulation of these zones revealed summation, reciprocity response interaction, or the absence of their interaction by the sensitivity index (ratio between response magnitudes to a cruciform figure and a bar). Stimulation of only central or only peripheral zone of the RF in nearly equal number of cases increased or decreased the sensitivity index in comparison with full cruciform figure. The neurons were found which did not change their sensitivity to cruciform figure under conditions of its partial masking. Mechanisms and functional significance of the interaction between the central and peripheral zones of RF for selection of line crossing in the striate cortex are discussed.

Animals↗

Postnatal development of NR1, NR2A and NR2B immunoreactivity in the visual cortex of the rat.

N-Methyl-D-aspartate receptors (NMDARs) are critically involved in some types of synaptic plasticity. The NMDAR subunits NR1, NR2A and NR2B are developmentally regulated, and it has been proposed that developmental changes in their expression may underlie developmental changes in cortical plasticity. Age-dependent change in cortical plasticity is most commonly measured by the monocular deprivation effect, which occurs during a critical period between P22 and P50 in the rat. Although the development of NMDAR subunits has been studied from birth through the fourth postnatal week, there is only meager information from older ages when visual plasticity ends. We hypothesized that there will be significant age-dependent change in expression of NR1, NR2A or NR2B between P22, when the cortex is plastic, and P90, when it is not. We applied specific antibodies recognizing NR1, NR2A and NR2B to the primary visual cortex at P14, P22, P30, P45 and P90. We found age-dependent changes in NR1-IR that were negatively correlated with changes in NR2A-IR; these subunits are not regulated in unison. In contrast, NR2A-IR and NR2B-IR were positively correlated. NR2A-IR and NR2B-IR both passed through a developmental minimum around P45, then recovered to approximately their P22 level. NR1-IR passed through a maximum at P45. There were no significant differences between P22 and P90. These results do not support the simple hypothesis that the loss of plasticity corresponds to a simple transition from juvenile levels of NMDAR subunit proteins to new adult levels. On the other hand, the results do confirm the hypothesis that there are significant changes in processing of NMDAR proteins during the time that plasticity is lost. How these changes of IR relate to synaptic transmission and plasticity needs to be clarified.

Animals↗

Functional MR of brain activity and perfusion in patients with chronic cortical stroke.

PURPOSE: (1) To determine whether functional MR can reliably map functional deficits in patients with stroke in the primary visual cortex; (2) to determine whether functional MR can reliably map perfusion deficits; and (3) to determine whether functional MR can give any additional diagnostic information beyond conventional MR. METHODS: Seven patients who had had a stroke in their primary visual system were examined using two functional MR techniques: (1) dynamic susceptibility contrast imaging, and (2) cortical activation mapping during full-field visual stimulation. Maps of relative cerebral blood volume and activation were created and compared with visual field examinations and conventional T2-weighted images on a quadrant-by-quadrant basis in five of these patients. RESULTS: Visual field mapping matched with both T2-weighted conventional images and activation mapping of 16 of 18 quadrants. In two quadrants, the activation maps detected abnormalities that were present on the visual field examination but not present on the T2-weighted images nor on the relative cerebral blood volume maps, which may indicate abnormal function without frank infarction. In addition, the activation maps demonstrated decreased activation in extrastriate cortex and had normal T2 signal and relative cerebral blood volume but was adjacent to infarcted primary cortex, mapping in vivo how stroke in one location can affect the function of distant tissue. CONCLUSION: Functional MR techniques can accurately map functional and perfusion deficits and thereby provide additional clinically useful information. Additional studies will be needed to determine the prognostic utility of functional MR in stroke patients.

Aged↗

The McCollough effect reveals orientation discrimination in a case of cortical blindness.

BACKGROUND: The McCollough effect is a colour after-effect that is contingent on the orientation of the patterns used to induce it. To produce the effect, two differently oriented grating patterns--such as a red-and-black vertical grating and a green-and-black horizontal grating--are viewed alternatively for a few minutes. After this period of adaptation, if the black-and-white test gratings are viewed in the same orientation as the adaptation patterns, the white sections of the vertical grating will appear pale green and the white sections of the horizontal grating will appear pink. The McCollough effect indicates that colour- and orientation-coding mechanisms interact at some point during visual processing; but the question remains as to whether this interaction occurs at an early or later stage in the cortical visual pathways. In an attempt to answer this question, we studied a patient who had suffered extensive damage to extrastriate visual areas of the brain, which had left him able to see colour but little else. RESULTS: Neuropsychological and perceptual tests demonstrated that the patient, P.B., has a profound impairment in form perception and is even unable to discriminate between 90 degrees differences in the orientation of grating stimuli. He is also unable to use orientation information to control his reaching or grasping. Nevertheless, P.B. can name and discriminate different colours reliably, including those used to induce the McCollough effect. After adaptation with red-and-green gratings, P.B. appropriately reported the orientation-contingent aftereffect colours, even though he continued to be unable to discriminate the orientations of the test patterns. CONCLUSIONS: These results indicate that at some level in P.B.'s visual system orientation is being coded, but it is at a level that he is unable to use in making orientation judgements or in visuomotor control. Given the massive insult to the extrastriate cortex in P.B., it is likely that the anatomical locus of the mechanisms underlying the McCollough effect is within primary visual cortex or even earlier in the visual pathway.

Adult↗

Differential quenching and limits of resolution in autoradiograms of brain tissue labeled with 3H-, 125I- and 14C-compounds.

Problems in interpretation of autoradiograms generated by ligand binding in brain tissue may be caused by two types of technical limitations: the differential absorption of 3H-generated emissions within the tissue (differential quenching) and the reduced resolution when 125I and 14C are used as isotopes. In the course of our ongoing receptor binding studies in primate brain, we have examined these methodological problems using neocortex of adult rhesus monkey as an example of a complex multilayered brain structure. We have compared: (1) film images produced by brain sections mounted on 3H- and 14C-labeled plastic; (2) autoradiograms of sections labeled with pairs of similar compounds containing 3H, 125I or 14C; and (3) autoradiograms of normal and defatted brain sections. The results indicate that differential absorption of 3H-generated emissions presents a genuine problem for film autoradiography of neocortex of adult monkey when 3H-compounds are used. Particularly significant attenuations of 3H-generated emissions are associated with sublayer IVb of primary visual cortex (Brodmann's are 17) and layers III (deep strata), V and VI of primary motor cortex (Brodmann's area 4). This study provides the necessary corrections for autoradiographic measurements. We also found a loss of resolution associated with use of 125I and 14C, a result that poses a significant problem for analysis of fine laminar patterns of the neocortex in adult monkeys. The use of isotopes with high energy emissions tends to decrease the variations in optical densities within the autoradiograms of cortical sections. Thus, the variations in optical density of autoradiograms of cortices labeled with 125I- and 14C-compounds may not represent the true distribution of these compounds.

Animals↗

Mechanisms of central color vision.

In monkey cerebral cortex, color information is processed along the ventral visual pathway. This pathway starts in the primary visual cortex and ends in area TE of the inferior temporal cortex. Recent studies indicate that the transformation of cone signals occurs early in the pathway to form neurons selective to a narrow range of hues. In addition, it has become apparent that area TE plays a vital role in color discrimination.

Animals↗

Analysis of fMRI data by blind separation of data in a tiny spatial domain into independent temporal component.

Independent Component Analysis (ICA) is a promising tool for the analysis of functional magnetic resonance imaging (fMRI) time series. In these studies, mostly assumed is a spatially independent component map of fMRI data (spatial ICA). In this paper, we assume that the temporal courses of the signal and noises are independent within a Tiny spatial domain (temporal ICA). Then with fast-ICA algorithm, spatially neighboring fMRI data were blindly separated into several temporal courses and were preassumed to be formed by a signal time course and several noise time courses where the signal has the largest correlation coefficient with the reference signal. The final functional imaging was completed for the signals obtained from each voxel. Simulations showed that compared with the spatial ICA method, the new temporal ICA method is more effective than the spatial ICA in detecting weak signal in a fMRI dataset. As background noise, the simulations include simulated Gaussian noise and fMRI data without stimulation. Finally, vivo fMRI tests showed that the excited areas evoked by a visual stimuli are mainly in the region of the primary visual cortex and that evoked by auditory stimuli are mainly in the region of the primary temporal cortex.

Algorithms↗

The organization of somatosensory cortex in the short-tailed opossum (Monodelphis domestica).

The organization of neocortex in the short-tailed opossum (Monodelphis domestica) was explored with multiunit microelectrode recordings from middle layers of cortex. Microelectrode maps were subsequently related to the chemoarchitecture of flattened cortical preparations, sectioned parallel to the cortical surface and processed for either cytochrome oxidase (CO) or NADPH-diaphorase (NADPHd) histochemistry. The recordings revealed the presence of at least two systematic representations of the contralateral body surface located in a continuous strip of cortex running from the rhinal sulcus to the medial wall. The primary somatosensory area (S1) was located medially while secondary somatosensory cortex (S2) formed a laterally located mirror image of S1. Auditory cortex was located in lateral cortex at the caudal border of S2, and some electrode penetrations in this area responded to both auditory and somatosensory stimulation. Auditory cortex was outlined by a dark oval visible in flattened brain sections. A large primary visual cortex (V1) was located at the caudal pole of cortex, and also consistently corresponded to a large chemoarchitecturally visible oval. Cortex just rostral and lateral to V1 responded to visual stimulation, while bimodal auditory/visual responses were obtained in an area between V1 and somatosensory cortex. The results are compared with brain organization in other marsupials and with placentals and the evolution of cortical areas in mammals is discussed.

Animals↗

Temporal analysis of the flow from V1 to the extrastriate cortex in humans.

We previously examined the cortical processing in response to somatosensory, auditory and noxious stimuli, using magnetoencephalography in humans. Here, we performed a similar analysis of the processing in the human visual cortex for comparative purposes. After flash stimuli applied to the right eye, activations were found in eight cortical areas: the left medial occipital area around the calcarine fissure (primary visual cortex, V1), the left dorsomedial area around the parietooccipital sulcus (DM), the ventral (MOv) and dorsal (MOd) parts of the middle occipital area of bilateral hemispheres, the left temporo-occipito-parietal cortex corresponding to human MT/V5 (hMT), and the ventral surface of the medial occipital area (VO) of the bilateral hemispheres. The mean onset latencies of each cortical activity were (in ms): 27.5 (V1), 31.8 (DM), 32.8 (left MOv), 32.2 (right MOv), 33.4 (left MOd), 32.3 (right MOv), 37.8 (hMT), 46.9 (left VO), and 46.4 (right VO). Therefore the cortico-cortical connection time of visual processing at the early stage was 4-6 ms, which is very similar to the time delay between sequential activations in somatosensory and auditory processing. In addition, the activities in V1, MOd, DM, and hMT showed a similar biphasic waveform with a reversal of polarity after 10 ms, which is a common activation profile of the cortical activity for somatosensory, auditory, and pain-evoked responses. These results suggest similar mechanisms of the serial cortico-cortical processing of sensory information among all sensory areas of the cortex.

Adult↗

[Adaptation of the central nervous system to optical correction].

Wearing spectacles imply an adjustment of the visual perception and eye movements. The visual cortex accounts for this plasticity, including at the adulthood, especially by the shift or the sprading of the receptor fields and the adjustment of the sensitivity of the primary visual cortex cells to spatial orientation and movement. The cerebellum modulates the vestibulo-ocular reflex gain. The adjustment latencies range from a few minutes to several days according to the disturbancy severity, the drug interferences and the age and medical history of the subject. Neurotrophins seem to be essential for this adjustment and might become an efficient tool to extend the plasticity period.

Adaptation, Physiological↗

Experience-dependent plasticity without long-term depression by type 2 metabotropic glutamate receptors in developing visual cortex.

Synaptic depression is thought to underlie the loss of cortical responsiveness to an eye deprived of vision. Here, we establish a fundamental role for type 2 metabotropic glutamate receptors (mGluR2) in long-term depression (LTD) of synaptic transmission within primary visual cortex. Direct mGluR2 activation by (2S,2'R,3'R-2-(2',3'-dicarboxycyclopropyl)glycine (DCG-IV) persistently depressed layer 2/3 field potentials in slices of mouse binocular zone when stimulated concomitantly. Chemical LTD was independent of N-methyl-d-aspartate (NMDA) receptors but occluded conventional LTD by low-frequency stimulation, indicating shared downstream events. Antagonists or targeted disruption of mGluR2 conversely prevented LTD induction by electrical low-frequency stimulation to layer 4. In contrast, Schaeffer collateral synapses did not exhibit chemical LTD, revealing hippocampal area CA1, naturally devoid of mGluR2, to be an inappropriate model for neocortical plasticity. Moreover, monocular deprivation remained effective in mice lacking mGluR2, and receptor expression levels were unchanged during the critical period in wild-type mice, indicating that experience-dependent plasticity is independent of LTD induction in visual cortex. Short-term depression that was unaffected by mGluR2 deletion may better reflect circuit refinement in vivo.

Animals↗

Magnetoencephalography reveals two distinct sources associated with late positive evoked potentials during visual oddball task.

The present investigation was undertaken in order to utilize magnetoencephalography to locate generator sources, modeled as equivalent current point dipoles, that account for the well-established late positive electrical potentials commonly measured along the midline (Cz, Pz, Fz, and Oz) in response to rare or task-relevant stimuli and their simultaneously recorded magnetic field components. Two simultaneous but spatially distinct sources were present in all six subjects. One source in which the magnetic flux exited and reentered the brain over the right hemisphere was localized in deep structures under the temporal cortex in the vicinity of the right hippocampal formation. The other occurred in the vicinity of the primary visual cortex in the occipital area with magnetic flux entering and exiting over the posterior aspects of the subjects' heads. These data correspond to depth and surface electrode studies that have demonstrated multiple generator sources.

Adult↗

Laminar and regional distributions of neurofibrillary tangles and neuritic plaques in Alzheimer's disease: a quantitative study of visual and auditory cortices.

The number of Thioflavine S-positive neurofibrillary tangles (NFT) and neuritic plaques (NP) was determined in visual and auditory cortical regions of 8 patients with Alzheimer's disease. On both a regional and laminar basis, NFT exhibited very distinctive and consistent distribution patterns. The mean (+/- SEM) number of NFT in a 250-micron-wide cortical traverse was very low in area 17, primary visual cortex (0.9 +/- 1.0), increased 20-fold in the immediately adjacent visual association cortex of area 18 (19.7 +/- 3.6), and showed a further doubling in area 20, the higher-order visual association cortex of the inferior temporal gyrus (35.5 +/- 8.8). Similar differences in NFT number were present between primary auditory (1.6 +/- 0.5) and auditory association (18.9 +/- 5.4) regions. On a laminar basis, NFT were predominantly present in layers III and V, although there were striking regional differences in the proportion of NFT in these 2 layers. Layer III contained 79% of the NFT in layers III and V in area 18, 41% in area 20, and only 27% in area 22. In contrast, NP showed different, and less specific, regional and laminar distribution patterns. Total NP number was similar in the 3 visual areas, although there were marked regional differences in the type of NP present. Nearly 80% of the NP in area 17 was of the NPc type (i.e., contained a dense, brightly fluorescent core), whereas over 70% of the NP in both areas 18 and 21 was of the NPnc type (i.e., lacked a dense, brightly fluorescent core). NP were present in every cortical layer but were most numerous in layers III and IV. The distinctive distribution patterns of NFT are very similar to the regional and laminar locations of long corticocortical projection neurons in homologous regions of monkey neocortex. This association suggests that NFT reside in the cell bodies of a subpopulation of pyramidal neurons, namely, those that furnish long corticocortical projections. In contrast, the distribution patterns of NP suggest that multiple neuronal systems contribute to their formation.

Aged↗

Effects of aging on the neuroglial cells and pericytes within area 17 of the rhesus monkey cerebral cortex.

An electron microscopic analysis has been carried out to compare the neuroglial cells and pericytes within the primary visual cortex, area 17, of young (5-6 years) and old (25-35 years) rhesus monkeys. All of the neuroglial cell types accumulate inclusions within their cytoplasm as they age, and the inclusions within the astrocytes and oligodendrocytes are essentially characteristic of those cell types. The astrocytes probably acquire their inclusions by phagocytosis, and it is suggested that the inclusions in the oligodendrocytes are caused by an age-related degeneration of the myelin sheaths they produce. The inclusions within the microglia are very heterogeneous. They are more massive than in the other neuroglial cells, so that their inclusions may almost fill the microglia. Pericytes also accumulate inclusions with age and there is evidence to suggest that they empty the contents of their inclusions vacuoles directly into the capillaries. On the basis of counts of the numbers of profiles of neuroglial cells displaying nuclei in thin sections, the only cells to increase in number with age are the microglia. They show an increase of about 44% when the cortices of young and old monkeys are compared.

Aging↗