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Surgically created neural pathways mediate visual pattern discrimination.

Combined lesions of retinal targets and ascending auditory pathways can induce, in developing animals, permanent retinal projections to auditory thalamic nuclei and to visual thalamic nuclei that normally receive little direct retinal input. Neurons in the auditory cortex of such animals have visual response properties that resemble those of neurons in the primary visual cortex of normal animals. Therefore, we investigated the behavioral function of the surgically induced retino-thalamo-cortical pathways. We showed that both surgically induced pathways can mediate visually guided behaviors whose normal substrate, the pathway from the retina to the primary visual cortex via the primary thalamic visual nucleus, is missing.

Animals↗

Emergent properties of layer 2/3 neurons reflect the collinear arrangement of horizontal connections in tree shrew visual cortex.

The superficial layers of primary visual cortex, unlike layer 4, have an extensive network of long-range horizontal connections linking sites of similar orientation preference. To identify possible functional consequences of this distinct anatomy, we compared the receptive field properties of layers 2/3 and 4 neurons in tree shrew primary visual cortex with electrophysiological recordings. We found that elongated receptive fields, strong orientation tuning, and length summation (properties predicted by the anatomy of the horizontal connections) are present in layer 2/3 neurons, but not in layer 4 neurons. We further characterized the summation fields of layer 2/3 neurons and found axis and orientation-specific facilitation that matched the distribution of horizontal connections. The functional signature of horizontal connections was also evident in the population response of layer 2/3 neurons; the intrinsic signal activation pattern elicited by an array of collinear Gabor elements was significantly stronger than that elicited by a noncollinear array. Furthermore, our results showed that this enhancement of population response was achieved without compromising spatial resolution along the collinear axis, providing stimulus-specific facilitation without filling in between stimuli. Taken together, these results suggest that horizontal connections play a significant role in shaping the visual responses of layer 2/3 neurons.

Animals↗

A functional MRI study of mental image generation.

The neural substrates of mental image generation were investigated with functional MRI. Subjects listened to words under two different instructional conditions: to generate visual mental images of the words' referents, or to simply listen to each word and wait for the next word. Analyses were performed which directly compared the regional brain activity during each condition, with the goal of discovering whether mental image generation engages modality-specific visual areas, whether it engages primary visual cortex, and whether it recruits the left hemisphere to a greater extent than the right. Results revealed that visual association cortex, and not primary visual cortex, was engaged during the mental image generation condition. Left inferior temporal lobe (Brodmann's area 37) was the most reliably and robustly activated area across subjects, had activity which extended superiorly into occipital association cortex (area 19). The results of this experiment support the hypothesis that visual mental imagery is a function of visual association cortex, and that image generation is asymmetrically localized to the left.

Adolescent↗

Representation of spatial frequency and orientation in the visual cortex.

Knowledge of the response of the primary visual cortex to the various spatial frequencies and orientations in the visual scene should help us understand the principles by which the brain recognizes patterns. Current information about the cortical layout of spatial frequency response is still incomplete because of difficulties in recording and interpreting adequate data. Here, we report results from a study of the cat primary visual cortex in which we employed a new image-analysis method that allows improved separation of signal from noise and that we used to examine the neurooptical response of the primary visual cortex to drifting sine gratings over a range of orientations and spatial frequencies. We found that (i) the optical responses to all orientations and spatial frequencies were well approximated by weighted sums of only two pairs of basis pictures, one pair for orientation and a different pair for spatial frequency; (ii) the weightings of the two pictures in each pair were approximately in quadrature (1/4 cycle apart); and (iii) our spatial frequency data revealed a cortical map that continuously assigns different optimal spatial frequency responses to different cortical locations over the entire spatial frequency range.

Animals↗

Visual projections routed to the auditory pathway in ferrets: receptive fields of visual neurons in primary auditory cortex.

How does cortex that normally processes inputs from one sensory modality respond when provided with input from a different modality? We have addressed such a question with an experimental preparation in which retinal input is routed to the auditory pathway in ferrets. Following neonatal surgical manipulations, a specific population of retinal ganglion cells is induced to innervate the auditory thalamus and provides visual input to cells in auditory cortex (Sur et al., 1988). We have now examined in detail the visual response properties of single cells in primary auditory cortex (A1) of these rewired animals and compared the responses to those in primary visual cortex (V1) of normal animals. Cells in A1 of rewired animals differed from cells in normal V1: they exhibited larger receptive field sizes and poorer visual responsivity, and responded with longer latencies to electrical stimulation of their inputs. However, striking similarities were also found. Like cells in normal V1, A1 cells in rewired animals exhibited orientation and direction selectivity and had simple and complex receptive field organizations. Furthermore, the degree of orientation and directional selectivity as well as the proportions of simple, complex, and nonoriented cells found in A1 and V1 were very similar. These results have significant implications for possible commonalities in intracortical processing circuits between sensory cortices, and for the role of inputs in specifying intracortical circuitry.

Animals↗

Feedback training of 36 - 44 HZ EEG activity in the visual cortex and hippocampus of cats: evidence for sensory and motor involvement.

Milk reinforcement was contingent on the occurrence of 36 -44 (40) Hz EEG activity in the left visual cortex (VC) of one group of cats and in the right hippocampus (H) of a second group. Both groups learned to increase 40 Hz activity, and acquisition of reinforcement was associated with immobility. A third group (behavioral controls - BC) was trained by the method of successive approximation to behave in a similar manner to VC and H cats. Training significantly increased 40 Hz activity in all of the following structures, except the hippocampi of VC cats and between the right and left visual cortex of H and BC cats: posterior primary visual cortex (bilateral), anterior primary visual cortex (left), primary motor cortex (bilateral), dorsal hippocampus (bilateral), and midbrain reticular formation (bilateral). Since the behavioral and EEG changes of H and BC animals were similar, immobility appears to be important for increased hippocampal 40 Hz activity produced by feedback training. Testing in darkness enhanced 40 Hz activity in the trained area of VC cats but had no effect on H or BC animals. These results, in conjunction with the observation that VC cats appeared to visually fixate, suggest that VC cats may have learned to increase 40 Hz activity in the visual cortex by altering visual processing.

Animals↗

Arrangement of ocular dominance columns in human visual cortex.

The arrangement of the ocular dominance columns in the human primary visual cortex was studied by examining cytochrome oxidase activity in autopsy specimens of occipital lobes obtained from two patients who became blind in one eye before death. By artificially flattening the cortex before processing, it was possible to reconstruct the pattern formed by the ocular dominance columns throughout most of the primary visual cortex. The columns form a mosaic of irregular parallel stripes about 500 microns to 1000 microns wide (right eye column plus left eye column measures 1 to 2 mm), oriented at right angles to the boundary of the primary visual cortex. The columns are wider near the boundary of the primary visual cortex and within the representation of the peripheral visual field, the ocular dominance columns of the ipsilateral eye become fragmented until they disappear altogether at the border of the monocular crescent representation. The arrangement of ocular dominance columns in the human visual cortex is very similar to the pattern reported in the macaque monkey, although the columns in humans are wider.

Aged↗

A polysynaptic feedback circuit in rat visual cortex.

Feedback connections from extrastriate cortex to primary visual cortex (V1) in the primate may provide "top-down" information that plays a role in visual attention and object recognition. Our work in a rodent model of corticocortical circuitry demonstrates that feedback pathways synapse preferentially with pyramidal cells in V1 () and favor excitation over inhibition in cortical microcircuits (). To investigate the polysynaptic circuits activated by feedback inputs, we studied chains of neurons postsynaptic to feedback connections using a combination of axonal tract tracing and anterograde degeneration. This approach enabled independent labeling of local collaterals of forward-projecting neurons in V1 and feedback connections from extrastriate lateromedial (LM) visual area to V1. Postsynaptic targets were identified in the electron microscope after retrograde transport of biotinylated dextran amine (BDA) to identify dendrites of forward-projecting neurons (i.e., from V1 to LM) and postembedding immunogold labeling to identify GABAergic interneurons. The results show that feedback connections provide strong monosynaptic input to forward-projecting neurons in V1. These neurons in turn make local connections that preferentially form synapses with other pyramidal cells ( approximately 97%), many of which were identified as forward-projecting neurons. This indicates that feedback pathways provide input directly to neurons which make the reciprocal forward connection, and that feedback-recipient forward-projecting neurons are strongly interconnected. The function of these excitatory networks within V1 may be to amplify feedback activity and provide a circuit for modulation of striate cortical activity by top-down influences.

Animals↗

Interocular transfer of receptive field expansion in cat visual cortex.

Receptive fields in primary visual cortex have been shown to be capable of rapid expansion and contraction when exposed to an artificial scotoma, a masked segment of the visual field. To distinguish cortical from thalamic contributions to receptive field mutability, we tested interocular transfer of the effect in binocular cortical receptive fields, presenting the conditioning stimulus to the field in one eye and measuring size changes in the receptive field of the other eye. The expansion of the receptive fields in the non-conditioned eye was comparable to that in the conditioned eye. This result suggests that the expansion is due to mechanisms intrinsic to the cortex.

Animals↗

Monocular focal retinal lesions induce short-term topographic plasticity in adult cat visual cortex.

Electrophysiological recording in primary visual cortex (VI) was performed both prior to and in the hours immediately following the creation of a discrete retinal lesion in one eye with an argon laser. Lesion projection zones (LPZs; 21-64 mm2) were defined in the visual cortex by mapping the extent of the lesion onto the topographic representation in cortex. There was no effect on neuronal responses to the unlesioned eye or on its topographic representation. However, within hours of producing the retinal lesion, receptive fields obtained from stimulation of the lesioned eye were displaced onto areas surrounding the scotoma and were enlarged compared with the corresponding field obtained through the normal eye. The proportion of such responsive recording sites increased during the experiment such that 8-11 hours post-lesion, 56% of recording sites displayed neurons responsive to the lesioned eye. This is an equivalent proportion to that previously reported with long-term recovery (three weeks to three months). Responsive neurons were evident as far as 2.5 mm inside the border of the LPZ. The reorganization of the lesioned eye representation produced binocular disparities as great as 15 degrees, suggesting interactions between sites in VI up to 5.5 mm apart.

Animals↗

Summation and division by neurons in primate visual cortex.

Recordings from monkey primary visual cortex (V1) were used to test a model for the visually driven responses of simple cells. According to the model, simple cells compute a linear sum of the responses of lateral geniculate nucleus (LGN) neurons. In addition, each simple cell's linear response is divided by the pooled activity of a large number of other simple cells. The cell membrane performs both operations; synaptic currents are summed and then divided by the total membrane conductance. Current and conductance are decoupled (by a complementary arrangement of excitation and inhibition) so that current depends only on the LGN inputs and conductance depends only on the cortical inputs. Closed form expressions were derived for fitting and interpreting physiological data. The model accurately predicted responses to drifting grating stimuli of various contrasts, orientations, and spatiotemporal frequencies.

Animals↗

Spatial frequency maps in cat visual cortex.

Neurons in the primary visual cortex (V1) respond preferentially to stimuli with distinct orientations and spatial frequencies. Although the organization of orientation selectivity has been thoroughly described, the arrangement of spatial frequency (SF) preference in V1 is controversial. Several layouts have been suggested, including laminar, columnar, clustered, pinwheel, and binary (high and low SF domains). We have reexamined the cortical organization of SF preference by imaging intrinsic cortical signals induced by stimuli of various orientations and SFs. SF preference maps, produced from optimally oriented stimuli, were verified using targeted microelectrode recordings. We found that a wide range of SFs is represented independently and mostly continuously within V1. Domains with SF preferences at the extremes of the SF continuum were separated by no more than (3/4) mm (conforming to the hypercolumn description of cortical organization) and were often found at pinwheel center singularities in the cortical map of orientation preference. The organization of cortical maps permits nearly all combinations of orientation and SF preference to be represented in V1, and the overall arrangement of SF preference in V1 suggests that SF-specific adaptation effects, found in psychophysical experiments, may be explained by local interactions within a given SF domain. By reanalyzing our data using a different definition of SF preference than is used in electrophysiological and psychophysical studies, we can reproduce the different SF organizations suggested by earlier studies.

Algorithms↗

Functional retinotopy of monkey visual cortex.

The operations of primary visual cortex generate continuous representations of orientation, ocular dominance, and retinotopy that, to fit in two dimensions, organize at separate but overlapping scales (e.g., 20-500 microm, 200 microm to 5 mm, and 2-33 mm). Where their scales overlap, these organizations interact; iso-orientation contours cross ocular dominance columns at right angles, and ocular dominance columns distort retinotopy near the V1/V2 border. To explore these interactions, we developed an optical technique for visualizing retinotopy in vivo that allows us to analyze it in relation to ocular dominance and orientation patterns. Our results show local retinotopic distortions in every region of macaque V1 that we examine, including regions far from the V1/V2 border. They also show a consistent relation between local axes of distortion and ocular dominance slabs, which they intersect at angles of approximately 90 degrees. A further correlation is provided by retinotopic maps from New World primates that show less distortion (9 vs 60%) in two species characterized by an absence of pronounced ocular dominance columns. Retinotopic maps from these New World primates also revealed an unexpected tilt of the vertical midline representation that diverged from the V1/V2 border by an angle of approximately 20 degrees. Overall, these results suggest a general tendency for slab-based organizations to distort retinotopy by representing the same part of space more than once in adjacent slabs.

Animals↗

Specialized vascularization of the primate visual cortex.

We have analyzed blood vessel distribution in the primary and secondary visual cortices of the squirrel monkey in relation to cortical modules, laminae, and cytoarchitectonic areas. Measurements of microvessel length in tangential sections through the primary visual cortex showed that blobs are more richly vascularized than intervening cortical regions. Thus, the mean total length of microvessel profiles per unit was 42% greater within these cortical modules than within adjacent (interblob) areas. Total microvessel length per unit area in another class of module, the stripes in the secondary visual cortex, was 27% greater than in interstripe regions. Microvessel distribution also varied systematically from layer to layer in the primary visual cortex, being greatest in lamina IVc. Finally, the overall microvessel length per unit area in sections of the primary visual cortex was 26% greater than that in the secondary visual cortex. These observations indicate that the modular, laminar, and regional organization of the primate visual cortex is reflected in the underlying distribution of cortical microvessels. These vascular patterns should be discernable in living animals with vascular contrast agents and appropriate imaging techniques.

Animals↗

Cortical activity blockade prevents ocular dominance plasticity in the kitten visual cortex.

Recordings from single units in kitten primary visual cortex show that a reversible blockade of the discharge activities of cortical neurons and geniculocortical afferent terminals by intracortical infusion of the sodium channel blocker tetrodotoxin (TTX) completely prevented the ocular dominance shift that would normally be seen after monocular deprivation. The blockade of cortical plasticity, like the blockade of discharge activity, was reversible, and plasticity was restored following recovery from the effects of TTX. These results extend previous work suggesting involvement of electrical activity at the level of the cortex in the phenomenon of cortical plasticity by demonstrating an absolute requirement for discharge activities in the primary visual cortex.

Action Potentials↗

Functional MRI of brain activation by eye blinking.

Functional magnetic resonance imaging (fMRI) was used to map cortical areas that control eye blinking. T2*-weighted asymmetric spin-echo MRI (or BOLD-blood oxygen level dependent-MRI) was used to detect changes caused by focal variations in blood oxygenation. Six normal volunteers and two cases of dry eye (less than 5-mm Schirmer's test) entered the study. The experimental scheme consisted of three cycles of a two-step sequence: (eyes closed)-(blink or blink inhibition). And to minimize contamination from photic activation, the experiments were carried out in a dark environment and the volunteers reported no light perception during the MR scans. In all eight cases, normal blinking (about one blink every 4 sec) activated areas in the orbitofrontal cortex and in some cases, the visual cortex including the anterior portion of the visual cortex and the primary visual cortex. In severe dry eye, blink-inhibition strongly activated the visual cortex even after irritation due to corneal desiccation was removed by topical anesthesia. The blinking process, especially the rate, appears to be controlled in the orbitofrontal cortex. The significance of visual cortex activation in the dark and in the case of severe dry eye still remains unclear; although it may be associated with attention and arousal.

Adult↗

Imagery without perception--a case study of anosognosia for cortical blindness.

A patient with complete cortical blindness after bilateral posterior cerebral artery infarctions denied her blindness. Her pretended visual experiences could frequently be traced back to synaesthetic translations of acoustic or tactile perceptions into mental visual images. Possibly, the belief to see resulted from a confusion of mental visual images with real percepts. The patient manifested preserved visual imagery also by correct responses to questions concerning the shapes of letters and the shapes and colours of objects. MRI showed an almost complete destruction of primary visual cortex with sparing of only small remainders of cortex at the occipital tip of the left upper calcarine lip. In the literature there are a few cases of denial of blindness with similarly severe damage to primary visual cortex but none with unequivocal evidence of complete destruction of primary visual cortex. We conclude that severe damage to primary visual cortex is compatible with visual imagery but that there is a possibility that islands of visual cortex must be spared to permit the generation of mental visual images.

Blindness↗

Retinotopic organization of striate and extrastriate visual cortex in the mouse.

Detailed retinotopic maps of primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18b) have been constructed for the C57BL/6J mouse using standard electrophysiological mapping techniques. Primary visual cortex (area 17), as defined cytoarchitectonically, contains one complete representation of the contralateral visual field, termed V1, in which azimuth and elevation lines are approximately orthogonal. The upper visual field is represented caudally and the nasal field laterally. Binocular cells are encountered in the cortical representation of the nasal 30--40 degrees of the visual field, and there is an expanded representation of the nasal field. Extrastriate visual cortex of the mouse, like that of other mammals, contains multiple representations of the visual field. The cytoarchitectonic region of cortex lateral and rostral to area 17, termed area 18a, contains at least two such representations. The more medial of these, which by convention we have called V2, is a narrow strip surrounding V1 on its lateral and rostral aspects; the vertical meridian lies along a portion of its common border with V1. The visual field representation in V2 is not a mirror image of that in V1; the representation of the horizontal meridian forms the lateral border of V2, and the visual field representation is split so that adjacent points on either side of the horizontal meridian are represented in nonadjacent parts of V2. The other visual field representation within area 18a, which we have termed V3, is a small but apparently complete representation that lies lateral to V2. The visual field representations medial to area 17 correspond to cytoarchitectonic area 18b. Area 18b contains two representations of the temporal visual field that we have labeled Vm-r and Vm-c, and contains little or no representation of the most nasal aspect of the field.

Animals↗