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Eye dominance in the visual cortex using functional MRI at 1.5 T: an alternative method.

PURPOSE: To develop a functional MRI method for producing eye dominance histograms in humans at 1.5 Tesla (T). METHODS: In the first set of experiments, 8 normal persons were tested. The eye dominance of each voxel within the person's visually activated primary visual cortex was determined with Student t statistics during a left eye versus right eye contrast. Eye dominance distribution was plotted, and the mean t statistic was used to describe the histogram asymmetry. In the second set of experiments, the effect of monocular optical blur and decreased luminance via filter was studied, and eye dominance distributions were similarly determined. RESULTS: The eye dominance histogram in each of the 8 normals was approximately symmetric; the average mean t value was +0.13. All 4 subjects with the right eye blurred had histograms approximately symmetric or slightly shifted toward the left eye (average mean t = +0.56), and all 4 subjects with the right eye filtered had histograms dramatically shifted toward the left eye (average mean t = +2.22). The average mean t for the group with the right eye filtered was significantly different from that of the other 2 groups (P <.0001). CONCLUSIONS: With noninvasive methods in normal persons, functional magnetic resonance imaging techniques at 1.5 T were able to characterize the distribution of eye dominance of voxels in primary visual cortex, based upon their t statistic in the left eye versus right eye contrast. The method is sensitive to filtering but relatively insensitive to visual blur. This approach may have a future use in the study of amblyopia in humans.

Adolescent↗

[Connections between cat brain visual cortex fields 18 and 19 passing through the white matter].

Distribution of degenerating synaptic boutons in fields 18 and 19 of the cat cerebral visual cortex was studied in every layer electron microscopically after the white substance was dissected under one of these fields. It was demonstrated that the V-shaped system under investigation consisting of associative fibres connects four superficial cortical layers (in layers V and VI of fields 18 and 19 no altered synapses were detected), while the intracortical system of conductors unites all six layers of the fields mentioned. The number of synapses, degenerating ones including, terminating on large and small dendrites and on thorns of every layer of fields 18 and 19 were counted. Possible functional differences between intracortical and V-shaped systems of the associative fibres in fields 18 and 19 of the cat visual cortex is discussed.

Animals↗

[Detector neurons of the visual cortex of the chipmunk].

Three functional classes of neurons are described in the Siberian chipmunk visual cortex: neurons nonselective to movement direction, movement-direction-selective neurons and orientation-selective neurons. Nonselective and direction-selective neurons showed maximum adaptiveness at high speeds of movement: 100-500 degrees/s and more. Most of orientation-selective neurons were maximally activated at speeds 10-50 degrees/s. For all neuronal classes a clear-cut correlation between selectivity and movement speed as well as pattern of responses to stationary stimuli in the receptive field were found. The data obtained allowed a division of the neurons into two groups: phasically-fast and tonically-slow with predominance of the first group.

Animals↗

[Analysis of the stability of rabbit visual cortex neuron spike responses to light flashes].

The stability of neuronal responses in the rabbit visual cortex to diffuse photic flashes presented in series with sufficiently long time intervals between them was studied in acute experiments. Three types of neuronal responses with different degree of stability were singled out. The stability was judged by the mean coefficient of correlation between post-stimulus histograms of spike activity. Negative correlations were found between the mean correlation coefficient and mean square deviations of latencies of primary and secondary activation and the pause in impulse activity in neuronal responses to a photic flash. Positive correlations between primary activation latencies and mean square deviations of other response parameters suggest a greater stability of responses with a short latency.

Action Potentials↗

Patterns of synaptic input on corticocortical and corticothalamic cells in the cat visual cortex. I. The cell body.

Immunocytochemical and electron microscopic methods were used to examine the ultrastructure and synaptology of callosal and corticothalamic pyramidal cell somata in the cat visual cortex (area 17). Callosal and corticothalamic cells were labeled after injection of horseradish peroxidase (HRP) in the contralateral visual cortex or in the ipsilateral lateral geniculate nucleus. The synaptic relationship between each of the two populations of pyramidal cells and cells containing the inhibitory neurotransmitter gamma-aminobutyric acid (GABA) was examined at the light and electron microscope level using the combined techniques of retrograde transport of HRP and GABA immunocytochemistry. We found that callosal and corticothalamic cell somata have an ultrastructure and synaptology that distinguishes them from each other. Reconstructions from electron micrographs of serial sections revealed that the vast majority of synapses (89-96%) on the cell body of pyramidal cells were formed by GABAergic axon terminals, and that within each population of pyramidal cells there was variability in the number and density of axosomatic synapses. Callosal pyramidal cells received a greater number and higher density of axosomatic synapses than corticothalamic cells. These data suggest that callosal cells receive more inhibition than corticothalamic cells at the level of their somata.

Animals↗

Spontaneous activity in developing ferret visual cortex in vivo.

Multi-electrode extracellular recordings in area 17 of awake behaving ferrets were conducted to characterize the pattern of spontaneous activity in the developing visual cortex before eye opening. A linear array of 16 microwire electrodes was used to record extracellular neuronal activity across a 3.2 mm strip of visual cortex between postnatal days 22 and 28. Whereas synchronous bursts of activity were observed at all recording sites, cross-correlation analysis revealed that the timing of spike activity at all electrodes was not precisely correlated. Correlated activity between cortical sites exhibited a patchy organization having long-range components. Long-range correlated activity was observed between cortical patches that were separated by a mean distance of 1 mm. The spatial pattern of correlated activity persisted during transient lateral geniculate nucleus (LGN) activity block, indicating that long-range correlated activity is generated by intrinsic circuits within the cortex, independent of LGN input activity. These results demonstrate an innate patchy organization of correlated spontaneous activity within the cortex during the early development of cortical functional and anatomical organization.

Action Potentials↗

Independent component analysis of temporal sequences subject to constraints by lateral geniculate nucleus inputs yields all the three major cell types of the primary visual cortex.

Information maximization has long been suggested as the underlying coding strategy of the primary visual cortex (V1). Grouping image sequences into blocks has been shown by others to improve agreement between experiments and theory. We have studied the effect of temporal convolution on the formation of spatiotemporal filters--that is, the analogues of receptive fields--since this temporal feature is characteristic to the response function of lagged and non-lagged cells of the lateral geniculate nucleus. Concatenated input sequences were used to learn the linear transformation that maximizes the information transfer. Learning was accomplished by means of principal component analysis and independent component analysis. Properties of the emerging spatiotemporal filters closely resemble the three major types of V1 cells: simple cells with separable receptive field, simple cells with non-separable receptive field, and complex cells.

Action Potentials↗

A model for encoding multiple object motions and self-motion in area MST of primate visual cortex.

Many cells in the dorsal part of the medial superior temporal (MST) region of visual cortex respond selectively to specific combinations of expansion/contraction, translation, and rotation motions. Previous investigators have suggested that these cells may respond selectively to the flow fields generated by self-motion of an observer. These patterns can also be generated by the relative motion between an observer and a particular object. We explored a neurally constrained model based on the hypothesis that neurons in MST partially segment the motion fields generated by several independently moving objects. Inputs to the model were generated from sequences of ray-traced images that simulated realistic motion situations, combining observer motion, eye movements, and independent object motions. The input representation was based on the response properties of neurons in the middle temporal area (MT), which provides the primary input to area MST. After applying an unsupervised optimization technique, the units became tuned to patterns signaling coherent motion, matching many of the known properties of MST cells. The results of this model are consistent with recent studies indicating that MST cells primarily encode information concerning the relative three-dimensional motion between objects and the observer.

Animals↗

[Receptive fields of cat visual cortex neurons when photic stimulation parameters are altered].

Spatial excitability reliefs of receptive fields of visual cortex units were investigated in acute experiments on immobilized cats by changing physical and physiological parameters of light stimulation under dark, mesopic and low photopic adaptation. Transition from dark to low mesopic adaptation results in no changes of the shape and size of the receptive fields, revealed by constant intensity test slits, whereas with transition to low photopic adaptation the receptive fields were narrowed in 72% of cases and acquired an elongated shape while the sharpness of their selectivity to direction of movement and orientation of stimuli increases. At a constant background the receptive fields narrowed with decreasing brightness of the test light slit. The excitability relief in the field estimated by the absolute threshold of the cell response and by the level of intensity needed to get a constant number of discharges in response, considerably narrowed with a rise in the threshold criterion and with transition from dark to light adaptation. At light adaptation, stimulation with physiologically equal intensity revealed narrower reactivity reliefs. than those at dark adaptation. The estimation of the receptive fields, taking into account a possible contribution of light scattering in the eye media and over the screen, shows that in most cases their size cannot be explained by this factor.

Animals↗

Synaptic physiology of the flow of information in the cat's visual cortex in vivo.

Each stage of the striate cortical circuit extracts novel information about the visual environment. We asked if this analytic process reflected laminar variations in synaptic physiology by making whole-cell recording with dye-filled electrodes from the cat's visual cortex and thalamus; the stimuli were flashed spots. Thalamic afferents terminate in layer 4, which contains two types of cell, simple and complex, distinguished by the spatial structure of the receptive field. Previously, we had found that the postsynaptic and spike responses of simple cells reliably followed the time course of flash-evoked thalamic activity. Here we report that complex cells in layer 4 (or cells intermediate between simple and complex) similarly reprised thalamic activity (response/trial, 99 +/- 1.9 %; response duration 159 +/- 57 ms; latency 25 +/- 4 ms; average +/- standard deviation; n = 7). Thus, all cells in layer 4 share a common synaptic physiology that allows secure integration of thalamic input. By contrast, at the second cortical stage (layer 2+3), where layer 4 directs its output, postsynaptic responses did not track simple patterns of antecedent activity. Typical responses to the static stimulus were intermittent and brief (response/trial, 31 +/- 40 %; response duration 72 +/- 60 ms, latency 39 +/- 7 ms; n = 11). Only richer stimuli like those including motion evoked reliable responses. All told, the second level of cortical processing differs markedly from the first. At that later stage, ascending information seems strongly gated by connections between cortical neurons. Inputs must be combined in newly specified patterns to influence intracortical stages of processing.

Action Potentials↗

Analysis of object motion in the ventral part of the medial superior temporal area of the macaque visual cortex.

1. The medial superior temporal area (MST) is an extrastriate area of the macaque visual cortex. Cells in MST have large receptive fields and respond to moving stimuli with directional selectivity. We previously suggested that the dorsal part of MST is mainly involved in analysis of field motion caused by movements of the animal itself, because most cells in the dorsal part preferentially responded to movements of a wide textured field rather than to movements of a small stimulus. To determine whether the remaining ventral part of MST differs in function from the dorsal part, we examined properties of cells in the ventral part in comparison with those of cells in the dorsal part, using anesthetized and paralyzed preparation. 2. Most cells in the ventral part preferably responded to movements of a small stimulus rather than to movements of a wide textured field. 3. Although the cells in the ventral part did not respond to movements of a textured field over a large window, many of them began to respond when a small stationary object was introduced in front of the moving field. The direction to which the cells responded in this stimulus configuration was opposite to the direction in which they responded to movements of an object on a stationary background. Activities of these cells thus represented the direction of relative movement of an object on a background, irrespective of whether the image of the object or the background moved on the retina. 4. We conclude that the ventral part of MST is distinctive from the dorsal part of MST and is mainly involved in the analysis of object movements in external space.

Animals↗

Visual cortex of the dolphin: an image analysis study.

On cytoarchitectonic grounds we have identified two distinct types of cortical formations composing the lateral gyrus (visual cortex) of the dolphin and have termed these heterolaminar cortex and homolaminar cortex. The heterolaminar cortex occupies the medial and lateral banks of the entolateral sulcus whereas the homolaminar cortex occupies the remainder of the lateral gyrus both lateral and medial to the entolateral sulcus. Each of these cortices exhibits special cytoarchitectonic features, a major difference being that heterolaminar cortex contains an incipient layer IV whereas layer IV is clearly absent in homolaminar cortex. Quantitative imaging procedures reveal that there is greater laminar differentiation in heterolaminar than in homolaminar cortex. Golgi analysis of neuronal forms and dendritic architecture confirms this distinction between the two types of cortex composing the lateral gyrus. Computer-assisted morphometric methods have been applied to both types of cortex and indicate by a variety of parameters several quantitative differences in the cellular numbers, types, and organization in each type of cortex. Both types of cortex, homolaminar and heterolaminar, exhibit a markedly higher cellular density in the posterior sector of the lateral gyrus than in the anterior sector. We have also for the first time been able to identify a columnar type of organization of the cetacean visual cortex and have described two types of cytoarchitectonic columns, major and minor, in each of these types of cortex. Comparisons in organization of these basic columnar units between the bat, representing a prototypic brain, and the dolphin reveal many similarities but also major quantitative differences in type of organization between the visual cortices in these species. Marked differences are also seen between the cytoarchitectonic columnar organization of the visual cortices in the dolphin and columnar organization of striate cortex in the human brain, the number of columns per unit of cortex in the human being almost twice that seen in the dolphin brain. Some phylogenetic implications of these findings are discussed in relation to the so-called "initial" type of cortical organization reconstructed largely by retrospective inference.

Aging↗

Topographic organization of the orientation column system in large flat-mounts of the cat visual cortex: a 2-deoxyglucose study.

We developed a flat-mount technique in order to visualize, without additional reconstruction, the system of orientation columns in the cat visual cortex by using 2-deoxyglucose-autoradiography. Experimental animals were injected with 2-deoxyglucose and then stimulated for 45-60 minutes either with vertical or horizontal or oblique gratings alone or with vertical and horizontal gratings presented in alternation. In both areas 17 and 18 stimulation with either vertical or horizontal or oblique stripes produced similar and highly ordered patterns of parallel bands of increased 2-deoxyglucose uptake that were perpendicular to the boundaries of the areas. In area 17 they occasionally extended without interruption from the 17/18 border on the top of the lateral gyrus to the monocular segment in the splenial sulcus. Superposition of serial sections revealed that these bands were present in all cortical layers and in precise register along lines orthogonal to the lamination. The center-to-center spacing of the bands was 1.0-1.1 mm in area 17 and 1.2-1.4 mm in area 18. Stimulation with alternating vertical and horizontal contours led to a pattern the general organization of which resembled that induced by a single orientation but the spacing of which was reduced by a factor of 0.5. This strongly supports the concept that orientation is mapped in a system of parallel bands and argues against a recently formulated hypothesis that iso-orientation bands extend like spokes from centers that lack orientation selectivity (Braitenberg and Braitenberg, Biol. Cybern. 33:179-186, '79). Another characteristic feature, revealed by the flat-mount technique, was a periodic variation of 2-deoxyglucose uptake along the bands that gave them a beaded appearance. The mean center-to-center distance between adjacent beads on the same band was in the range of 0.9-1.2 mm and remained unchanged when horizontal and vertical gratings were presented in alternation. We propose that these beads reflect another columnar system whose features have yet to be determined.

Animals↗

Intrinsic connections of rat primary visual cortex: laminar organization of axonal projections.

The organization of local projections within the rat primary visual cortex (area 17) was investigated by tracing fibers with HRP in in vitro brain slices. The projections from different layers showed distinct laminar patterns. Layer 4 made a strong, topographically precise, projection to lower layer 2/3; weaker projections extended laterally and terminated diffusely in layer 2/3 but also ran vertically to layers 5 and 6. The connections of lower and upper layer 2/3 were reciprocal and point-to-point. Within layer 2/3, a large number of fibers ran horizontally and terminated at variable distances from the injection site without making terminal clusters. The main output from layer 2/3 was to layer 5. The most prominent projections from the upper half of layer 5 were to layers 2/3 and 6; lower layer 5, in contrast, made wide-ranging, clustered projections to layer 1, the bottom of layer 2/3, and the top of layers 4 and 5. The patches were 130-160 micron wide and spaced apart by 230-260 micron. The main projection that arose from the superficial layer 6 terminated in layer 4 above the injection site. In contrast, lower layer 6 made clustered projections to the layer 3/4 border, extending up to 2 mm in the coronal plane. The patches were 190-220 micron wide and spaced apart by 320-390 micron. Additional projections went to the layer 5/6 border and layers 1 and 2. These results indicate that geniculocortical input is processed through interlaminar connections that are topographically precise, widespread, or patchy. These connectivity patterns suggest a role for these connections in the transformation of functional maps between layers; focused projections preserve the architecture of the layers of origin, and diverging or patchy projections rearrange this organization and form new maps in the target layers (Lund: Annu. Rev. Neurosci. 11:253-288, '88). However, only a few interlaminar connections show one of these patterns in isolation, making it difficult to assign a single function to a particular connection. We, therefore, tentatively conclude that projections terminating in layers 1-4, with the possible exception of the connection between upper layer 6 and layer 4, transform functional maps. In contrast, the topographically precise projections from upper to lower layers preserve functional maps. The specific role of these connections in the construction of receptive field properties, however, is not known.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Rearrangement of synaptic connections with inhibitory neurons in developing mouse visual cortex.

Cortical inhibition is determined in part by the organization of synaptic inputs to gamma-aminobutyric acidergic (GABAergic) neurons. In adult rat visual cortex, feedforward (FF) and feedback (FB) connections that link lower with higher areas provide approximately 10% of inputs to parvalbumin (PV)-expressing GABAergic neurons and approximately 90% to non-GABAergic cells (Gonchar and Burkhalter [1999] J. Comp. Neurol. 406:346-360). Although the proportions of these targets are similar in both pathways, FF synapses prefer larger PV dendrites than FB synapses, which may result in stronger inhibition in the FF than in the FB pathway (Gonchar and Burkhalter [1999] J. Comp. Neurol. 406:346-360). To determine when during postnatal (P) development FF and FB inputs to PV and non-PV neurons acquire mature proportions, and whether the pathway-specific distributions of FF and FB inputs to PV dendrites develop from a similar pattern, we studied FF and FB connections between area 17 and the higher order lateromedial area (LM) in visual cortex of P15-42 mice. We found that the innervation ratio of PV and non-PV neurons is mature at P15. Furthermore, the size distributions of PV dendrites contacted by FF and FB synapses were similar at P15 but changed during the third to sixth postnatal weeks so that, by P36-42, FF inputs preferred thick dendrites and FB synapses favored thin PV dendrites. These results suggest that distinct FF and FB circuits develop after eye opening by rearranging the distribution of excitatory synaptic inputs on the dendritic tree of PV neurons. The purpose of this transformation may be to adjust differentially the strengths of inhibition in FF and FB circuits.

Aging↗

Influence of visual experience deprivation on the postnatal development of the microvascular bed in layer IV of the rat visual cortex.

Cerebral vascular density is correlated with metabolic demands, which increase in highly active brain areas. External inputs are an essential requirement in the modeling of the visual cortex. Experience-mediated development is very active during the first postnatal month, when congruous blood supply is needed. We studied the development of visual cortex vascularization in relation to experience, comparing rats raised in darkness with rats reared in normal conditions. Vascular density, vascular area and their ratio vs. neuronal density were calculated. Conventionally stained semi-thin sections were used to measure the vascular area by computer assisted morphometry. Animals from both groups were sampled at 14, 21, and 60 days postnatal (dpn). We found a significantly lower density of vessels and neurons as well as a smaller vascular area in dark-reared adult rats while no differences were founded at the other ages. Our results also show no differences between the ratio of vessels/neuron, and vascular area/neuron, between both groups. The absence of visual experience causes decrease of cortical activity which correlates with lower vessels density and vascular area, without their ratio/neuron being affected.

Age Factors↗

Effect of transcranial magnetic stimulation on single-unit activity in the cat primary visual cortex.

Transcranial magnetic stimulation (TMS) has become a well established procedure for testing and modulating the neuronal excitability of human brain areas, but relatively little is known about the cellular processes induced by this rather coarse stimulus. In a first attempt, we performed extracellular single-unit recordings in the primary visual cortex (area 17) of the anaesthetised and paralysed cat, with the stimulating magnetic field centred at the recording site (2 x 70 mm figure-of-eight coil). The effect of single biphasic TMS pulses, which induce a lateral-to-medial electric current within the occipital pole of the right hemisphere, was tested for spontaneous as well as visually evoked activity. For cat visual cortex we found that a single TMS pulse elicited distinct episodes of enhanced and suppressed activity: in general, a facilitation of activity was found during the first 500 ms, followed thereafter by a suppression of activity lasting up to a few seconds. Strong stimuli exceeding 50 % of maximal stimulator output could also lead to an early suppression of activity during the first 100-200 ms, followed by stronger (rebound) facilitation. Early suppression and facilitation of activity may be related to a more or less direct stimulation of inhibitory and excitatory interneurons, probably with different thresholds. The late, long-lasting suppression is more likely to be related to metabotropic or metabolic processes, or even vascular responses. The time course of facilitation/inhibition may provide clues regarding the action of repetitive TMS application.

Action Potentials↗

Spatial sampling by dendritic trees in visual cortex.

Kittens were reared in vertically or horizontally striped cylinders. After rearing exposures of 400-500 h, responses of single neurons were determined as a function of orientation of a square wave grating stimulus. These data suggest that the rearing environment did alter orientation preference in some of the kittens. The visual cortices of the stripe-reared kittens and of control kittens were impregnated according to a Golgi-Cox method. Dendrites of layer IV stellate cells were tracked and analyzed in three dimensions by a computer-microscope. Four methods of analyzing the spatial distribution of dendrites are described and discussed. Two methods previously described in the literature were not sufficiently sensitive to detect any differences among kittens exposed to vertical or horizontal stripes or to a control environment. Two newly developed methods were able to provide initial evidence for rearing effects on dendritic trees in visual cortex. The more detailed of these new methods describes the angular location of dendritic segments, with respect to standard brain axes, as a function of distance from the cell body. Data obtained by means of this method of dendritic angular distribution (DAD) plots suggests a number of conclusions. Rearing animals in a striped environment may influence the way in which dendrites of layer IV stellate cells of visual cortex distribute themselves in the neuropil. The effect of selective rearing on dendritic distribution does not appear to extend back to those portions of the dendritic tree closest to the cell body. This influence of rearing in a selective environment may be explained by hypothesizing that during development dendrites distribute themselves in ways that tend to maximize the effects of spatiotemporal summation for the postsynaptic neurons.

Animals↗