Orientation selectivity in rat primary visual cortex emerges earlier with low‐contrast and high‐luminance stimuli

In natural vision, rapid and sustained variations in luminance and contrast change the reliability of information available about a visual scene, and markedly affect both neuronal and behavioural responses. The hallmark property of neurons in primary visual cortex (V1), orientation selectivity, is unaffected by changes in stimulus contrast, but it remains unclear how sustained differences in mean luminance and contrast affect the time‐course of orientation selectivity, and the amount of information that neurons carry about orientation. We used reverse correlation with characterize the temporal dynamics of orientation selectivity in rat V1 neurons under four luminance‐contrast conditions. We show that orientation selectivity and mutual information between neuronal responses and stimulus orientation are invariant to contrast or mean luminance. Critically, the time‐course of the emergence of orientation selectivity was affected by both factors; response latencies were longer for low‐ than high‐luminance gratings, and surprisingly, response latencies were also longer for high‐ than low‐contrast gratings. Modelling suggests that luminance‐modulated changes in feedforward gain, in combination with hyperpolarization caused by high contrasts can account for our physiological data. The hyperpolarization at high contrasts may increase signal‐to‐noise ratios, whereas a more depolarized membrane may lead to greater sensitivity to weak stimuli.     

Quantitative morphologic classification of layer 5 neurons from mouse primary visual cortex

The understanding of any neural circuit requires the identification and characterization of all its components. Morphologic classifications of neurons are, therefore, of central importance to neuroscience. We use a quantitative method to classify neurons from layer 5 of mouse primary visual cortex, based on multidimensional clustering. To reconstruct neurons, we used Golgi impregnations and biocytin injections, as well as DiOlistics, a novel technique of labeling neurons with lipophilic dyes. We performed computerized 3-D reconstructions of 158 layer 5 cells to measure a series of morphologic variables. Principal component analysis and cluster analysis were used for the classification of cell types. Five major classes of cells were found: group 1 includes large pyramidal neurons with apical dendrites that reach layer 1 with an apical tuft; group 2 consists of short pyramidal neurons and large multipolar cells with "polarized" dendritic trees; group 3 is composed of less extensive pyramidal neurons; group 4 includes small cells; and group 5 includes another set of short pyramidal neurons in addition to "atypically oriented" cells. Our sample included a relatively homogeneous group of 27 neurons that project to the superior colliculus, which clustered mainly in group 1, thus supporting the validity of the classification. Cluster analysis of neuronal morphologies provides an objective method to quantitatively define different neuronal phenotypes and may serve as a basis for describing neocortical circuits.     

Modulation of neuronal responses in macaque primary visual cortex in a memory task

The primary visual cortex, a relatively early station in the visual pathway, has long been considered mainly as a site of basic feature detection but evidence is emerging that is consistent with the existence of feedback influences from higher cortical areas. We show that in a delayed match-to-sample memory task, where the monkey needs to remember both the visual pattern and its location, there is significant modulation of neuronal activity in the primary visual cortex suggestive of a feedback signal. Responses to identical patterns are remarkably different depending upon their place in the memory task. These modulatory influences are significantly less when the same visual patterns are shown during a simple fixation task, where these stimuli can be ignored and not attended to. The results indicate that neural processing specific to attentional and mnemonic functions can involve even primary sensory areas.     

Attentional load modifies early activity in human primary visual cortex

Recent theories of selective attention assume that the more attention is required by a task, the earlier are irrelevant stimuli filtered during perceptual processing. Previous functional MRI studies have demonstrated that primary visual cortex (V1) activation by peripheral distractors is reduced by higher task difficulty at fixation, but it remains unknown whether such changes affect initial processing in V1 or subsequent feedback. Here we manipulated attentional load at fixation while recording peripheral visual responses with high-density EEG in 28 healthy volunteers, which allowed us to track the exact time course of attention-related effects on V1. Our results show a modulation of the earliest component of the visual evoked potential (C1) as a function of attentional load. Additional topographic and source localization analyses corroborated this finding, with significant load-related differences observed throughout the first 100 ms post-stimulus. However, this effect was observed only when stimuli were presented in the upper visual field (VF), but not for symmetrical positions in the lower VF. Our findings demonstrate early filtering of irrelevant information under increased attentional demands, thus supporting models that assume a flexible mechanism of attentional selection, but reveal important functional asymmetries across the VF.     

Surround suppression sharpens orientation tuning in the cat primary visual cortex

In the primary visual cortex (V1), the response of a neuron to stimulation of its classical receptive field (CRF) is suppressed by concurrent stimulation of the extraclassical receptive field (ECRF), a phenomenon termed 'surround suppression'. It is also known that the orientation tuning of V1 neurons becomes sharper as the size of the stimulus increases beyond the CRF. However, there have been few quantitative investigations of the relationship between sharpening of orientation tuning and surround suppression. We examined this relationship in 73 V1 neurons recorded from anesthetized and paralysed cats using sinusoidal grating patches as stimuli. We found that sharpening of orientation tuning was significantly correlated with the strength of surround suppression for large stimuli that cover both CRF and ECRF. Furthermore, simulation analysis using a variety of tuning widths and most suppressive orientation of orientation-tuned surround suppression demonstrated that broadly orientation-tuned surround suppression sharpens orientation tuning for large gratings without shift in optimal orientation. Our findings suggest that one of the functional roles of surround suppression in V1 is enhancement of orientation discrimination for large and uniformly patterned objects.     

Independence of visuotopic representation and orientation map in the visual cortex of the cat

The representations of visual space and stimulus orientation were mapped in the cat primary visual cortex using electrophysiological recordings supplemented with intrinsic signal optical imaging. The majority of units displaced up to 600 micro m laterally had overlapping RFs both in orientation domains and around singularities of the orientation map. Quantitative comparison of these units revealed only a weak, positive correlation between the difference in their preferred orientations and RF separations (area 17: r = 0.09; area 18: r = 0.15). The occurrence of nonoverlapping RFs could be accounted for by random RF position scatter rather than by orientation difference between the units. Monte Carlo analysis showed that our findings are compatible with a locally smooth and linear representation of visual space that is not coupled to the representation of stimulus orientation. An important functional implication of the above map relationships is that positional information captured by the retina is faithfully transmitted into the cortex.     

Retinotopic distribution of chromatic responses in human primary visual cortex

In non-human primates at least three anatomically and functionally distinct channels convey signals from the retina to the primary visual cortex (V1). Two of these channels, the parvocellular and the koniocellular, are sensitive to chromatic contrasts and form the basis of color vision. In humans, common phylogenetic history with other primates and psychophysical experiments suggest identical retinocortical mechanisms but separate evaluation of the distinct anatomical channels has been difficult because signals are already combined in V1. We studied the spatial distribution of activation to chromatic stimuli along the two opponent chromatic axes in human V1 with multifocal functional magnetic resonance imaging. The signal strength was quantified from three experiments with stimuli up to 20 degrees eccentricity. The hypothesis was that, although the parvo- and koniocellular signals are mixed in V1, distinct distributions of signal strength would be evident. We found that whereas different conditions activated the same areas of cortex, indicating that they have identical magnification factors, the responses to red/green stimulation were stronger close to the fovea whereas the blue/yellow responses were much less diminished with increasing eccentricity. Both chromatic axes showed saturating contrast response functions. Our measure directly from human V1 is in line with earlier psychophysical studies suggesting relatively stronger parvocellular channel representation close to the fovea, and more uniform distribution of the koniocellular and achromatic channels. In addition, our study presents a way to rapidly quantify retinotopic signal transmission in distinct retinocortical pathways of individual subjects.     

Functional parcellation of the human primary somatosensory cortex to natural touch

Despite the significance of human touch, brain responses to interpersonal manual touch have been rarely investigated. We used functional magnetic resonance imaging to study brain activity in eight healthy adults whose left hand was touched by two individuals, in separate runs and in 20‐s blocks, either by holding, smoothing, or poking. Acceleration was measured from both the subject's and the touching person's hands for postimaging control of the stimuli. Independent component analysis of the functional magnetic resonance imaging data unraveled three functional networks involving the primary somatosensory cortex (SI). One network comprised the contralateral and another the ipsilateral Brodmann area 3. The third network included area 2 bilaterally, left‐hemisphere middle temporal gyrus and dorsolateral prefrontal regions, ventral prefrontal cortices bilaterally, and middle cingulate cortex. The response shapes and polarities varied between the three networks. The contralateral area 3 differentiated the responses between the three types of touch stimuli, and the response magnitudes depended on the variability of the touch within each block. However, the responses of the other two networks were strikingly similar to all stimuli. The subjects' reports on the pleasantness of the touch did not correlate with the characteristics of the SI responses. These findings imply area‐specific processing of the natural human touch in three networks including the SI cortex, with only area 2 connected to a functional network of brain areas that may support social interaction.     

Anisomycin activates p38 MAP kinase to induce LTD in mouse primary visual cortex

Anisomycin is both a well-established protein synthesis inhibitor and a potent activator of the p38/JNK MAPK pathway. It has been used to block the late phase of long-term potentiation (LTP) and long-term depression (LTD) in hippocampus. In this study, we have found that anisomycin produces a time-dependent decline in the magnitude of the field EPSP (fEPSP) in acute brain slices of mouse primary visual cortex. This anisomycin-mediated fEPSP depression occludes NMDA receptor-dependent LTD induced by low-frequency stimulation (LFS). In contrast, two other protein synthesis inhibitors, emetine and cycloheximide, have no effect either on baseline synaptic transmission or on LTD. Moreover, the decline of the fEPSP caused by anisomycin can be rescued by the application of the p38 inhibitor SB203580 but not by the JNK inhibitor SP600125. These results indicate that activation of p38 MAPK by anisomycin induces LTD and subsequently occludes electrically induced LTD. Also, the occlusion of LFS-LTD by anisomycin suggests that common mechanisms may be shared between the two forms of synaptic depression. Consistent with this view, bath application of a membrane permeant peptide derived from the carboxyl tail of GluR2 subunit of AMPA receptor, which specifically blocks regulated AMPA receptor endocytosis, thereby preventing the expression of LFS-induced LTD, significantly reduced the anisomycin-induced decline of the fEPSP. In conclusion, our results indicate that anisomycin produces long-lasting depression of AMPA receptor-mediated synaptic transmission by activating p38 MAPK-mediated endocytosis of APMA receptors in mouse primary visual cortex.     

Role of primary visual cortex in the binocular integration of plaid motion perception

This study assessed the early mechanisms underlying perception of plaid motion. Thus, two superimposed gratings drifting in a rightward direction composed plaid stimuli whose global motion direction was perceived as the vector sum of the two components. The first experiment was aimed at comparing the perception of plaid motion when both components were presented to both eyes (dioptic) or separately to each eye (dichoptic). When components of the patterns had identical spatial frequencies, coherent motion was correctly perceived under dioptic and dichoptic viewing condition. However, the perceived direction deviated from the predicted direction when spatial frequency differences were introduced between components in both conditions. The results suggest that motion integration follows similar rules for dioptic and dichoptic plaids even though performance under dichoptic viewing did not reach dioptic levels. In the second experiment, the role of early cortical areas in the processing of both plaids was examined. As convergence of monocular inputs is needed for dichoptic perception, we tested the hypothesis that primary visual cortex (V1) is required for dichoptic plaid processing by delivering repetitive transcranial magnetic stimulation to this area. Ten minutes of magnetic stimulation disrupted subsequent dichoptic perception for approximately 15 min, whereas no significant changes were observed for dioptic plaid perception. Taken together, these findings suggest that V1 is not crucial for the processing of dioptic plaids but it is necessary for the binocular integration underlying dichoptic plaid motion perception.     

Optically imaged maps of orientation preference in primary visual cortex of cats and ferrets

Feature maps in the cerebral cortex constitute orderly representations of response features created within the cortex; an example is the mapping of orientation-selective neurons in visual cortex. We have compared the properties of orientation maps in area 17 of cats and ferrets, obtained by optical imaging of intrinsic signals. Orientation maps in both species contain a quasi-periodic distribution of iso-orientation domains that are organized into a lattice of pinwheels. However, the spatial density of orientation domains and of pinwheels in ferret area 17 is nearly twice that in cat area 17. The ferret map also contains more discontinuities, or fractures, where orientation changes abruptly. The size of orientation domains scales with interdomain spacing, so that the ratio of the two is approximately the same in both species. Consistent with this finding, the orientation tuning width of individual pixels is similar in the two. The magnitude of orientation preference, however, is much lower in ferret compared to cat. The greater incidence of fractures in ferret appears to be due to proportionately greater overlap between domains of different orientations, particularly along fracture lines that link pinwheel centers. We hypothesize that a key determinant of orientation maps, the relationship between orientation domain size and spacing, expresses an anatomical link between sizes of thalamocortical arbors and horizontal intracortical connections in area 17. J. Comp. Neurol. 387:358–370, 1997. © 1997 Wiley-Liss, Inc.     

Segregation of feedforward and feedback projections in mouse visual cortex

Hierarchical organization is a common feature of mammalian neocortex. Neurons that send their axons from lower to higher areas of the hierarchy are referred to as "feedforward" (FF) neurons, whereas those projecting in the opposite direction are called "feedback" (FB) neurons. Anatomical, functional, and theoretical studies suggest that these different classes of projections play fundamentally different roles in perception. In primates, laminar differences in projection patterns often distinguish the two projection streams. In rodents, however, these differences are less clear, despite an established hierarchy of visual areas. Thus the rodent provides a strong test of the hypothesis that FF and FB neurons form distinct populations. We tested this hypothesis by injecting retrograde tracers into two different hierarchical levels of mouse visual cortex (area 17 and anterolateral area [AL]) and then determining the relative proportions of double-labeled FF and FB neurons in an area intermediate to them (lateromedial area [LM]). Despite finding singly labeled neurons densely intermingled with no laminar segregation, we found few double-labeled neurons (≈5% of each singly labeled population). We also examined the development of FF and FB connections. FF connections were present at the earliest timepoint we examined (postnatal day 2, P2), while FB connections were not detectable until P11. Our findings indicate that, even in cortices without laminar segregation of FF and FB neurons, the two projection systems are largely distinct at the neuronal level and also differ with respect to the timing of their axonal outgrowth.     

Changes in projection from locus coeruleus to lateral geniculate nucleus following ablation of visual cortex in adult rats

The visual cortex of adult rats was unilaterally ablated. A histofluorescence study revealed an increase of noradrenergic terminals in the lateral geniculate nucleus (LGN) ipsilateral to the decortication, confirming the previous report. Corresponding to this, the frequency for encountering neurons in the locus coeruleus (LC) activated antidromically from the LGN was increased. We suggest that LC neurons whose axon terminals were damaged by ablation of the visual cortex formed new axons or axon collaterals (pruning effect) in the LGN, thus contributing, at least partly, to the increase of noradrenergic terminals therein.     

Sex differences in response to red and blue light in human primary visual cortex: a bold fMRI study

Studies using a variety of investigative methods, including functional brain imaging and electroencephalography (EEG), have suggested that changes in central nervous system (CNS) dopamine function result in altered visual system processing. The discovery of abnormal retinal blue cone, but not red cone, electroretinogram in association with cocaine withdrawal and Parkinson's disease suggests that visual system response to blue light might be a marker for CNS dopamine tone. As there are numerous sex-related differences in central nervous system dopamine function, we predicted that blue and red light stimulation would produce sex-specific patterns of response in primary visual cortex when studied using the blood oxygen level dependent (BOLD) functional magnetic resonance imaging (fMRI) technique. We analyzed the BOLD response to red and blue light in male and female human volunteers (N=20). Red and blue light responses in primary visual cortex (V1) to stepped intensities of red and blue light were compared by sex for threshold to detectable BOLD signal increase and for stimulus intensity vs. BOLD signal response. Near threshold, males and females showed similar BOLD signal change to red light, but males showed a threefold greater increase (0.52%) to blue light stimulation when compared to females (0.14%). Log-linear regression modeling revealed that the slope coefficients for the red light stimulus intensity vs. signal change curve were not significantly different for males and females (z=0.995, P=0.320), whereas the slope coefficients for the blue light stimulus intensity vs. signal change curve were significantly larger in males (z=2.251, P=0.024). These findings support a sex and color-dependent differential pattern of primary visual cortical response to photic stimulation and suggest a method for assessing the influence of specific dopamine agonist/antagonist medications on visual function.     

Trial‐by‐trial reliability of responses in the primary visual cortex on binocular disparity depends on stimulus order

An association of the detrimental effect of monocular deprivation on binocular vision with reduced reliability of neuronal responses in the primary visual cortex has been shown on randomly presented binocular stimuli [V. Vorobyov et al. (2007) Eur J Neurosci. 26(12), 3553–3563]. To examine this effect on biologically relevant signals, binocular gratings of varying relative phase disparity were presented in sequential order, simulating motion, to 55 cats with various types of daily visual experience. During sequential stimulation, the proportions of ‘unstable’ cells (with phase differences exceeding 22.5 ° between peak binocular responses in two consecutive trials) were similar in cats with exclusively binocular experience and with short periods of daily monocular vision (≤ 3.25 h), in mixed binocular–monocular conditions. In contrast, random stimulation was characterized by a significantly enlarged population of ‘unstable’ cells in the latter. After a longer period of monocular vision (6.5 h) or exclusively discordant binocular experience (strabismus), sequential stimulation was accompanied by a significant increase of this population, whereas during randomized stimulation it was very similar to that in cats with short periods of daily monocular vision. Finally, there were no differences in populations of ‘unstable’ cells in cats with long monocular or strabismic vision and those with exclusive monocular experience during sequential stimulation, in contrast with a significant increase in the latter during randomized stimulation. I propose that the detrimental effect of abnormal binocular experience on binocular processing in the primary visual cortex is associated with a disruption of the mechanisms involved in both discrimination of binocular disparity signals and evaluation of their temporal profiles.     

Contrast independence of cardinal preference: stable oblique effect in orientation maps of ferret visual cortex

The oblique effect was first described as enhanced detection and discrimination of cardinal orientations compared with oblique orientations. Such biases in visual processing are believed to originate from a functional adaptation to environmental statistics dominated by cardinal contours. At the neuronal level, the oblique orientation effect corresponds to the numerical overrepresentation and narrower tuning bandwidths of cortical neurons representing the cardinal axes. The anisotropic distribution of orientation preferences over large cortical regions was revealed with optical imaging, providing further evidence for the cortical oblique effect in several mammalian species. Our present study explores whether the dominant representation of cardinal contours persists at different stimulus contrasts. Performing intrinsic optical imaging in the ferret visual cortex and presenting drifting gratings at various orientations and contrasts (100%, 30% and 10%), we found that the overrepresentation of vertical and horizontal contours was invariant across stimulus contrasts. In addition, the responses to cardinal orientations were also more robust and evoked larger modulation depths than responses to oblique orientations. We conclude that orientation maps remain constant across the full range of contrast levels down to detection thresholds. Thus, a stable layout of the functional architecture dedicated to processing oriented edges seems to reflect a fundamental coding strategy of the early visual cortex.     

‘Real-motion’ cells in the primary visual cortex of macaque monkeys

Extracellular recordings were carried out in the primary visual cortex of behaving macaque monkeys. Neurons were activated by moving a visual stimulus across their receptive fields during periods of steady fixation and by moving their receptive fields (by visual tracking) across a motionless visual stimulus, taking care that the velocities of stimulus and eye movements were the same. The total cell population (108 neurons) ws divided into 3 groups according to the cell sensitivity to visual stimulus orientation (non-oriented cell and oriented cells) and to the presence or absence of antagonistic areas in in the receptive fields (oriented cells with antagonistic areas). All the non-oriented cells (n = 14) showed almost the same response to visual stimulation both during steady fixation and during visual tracking. Out of a total number of 86 oriented cells, 77 turned out to be activated by the visual stimulation both during fixation and tracking. Eight oriented cells gave a very weak response or no response at all to visual stimulation during smooth pursuit eye movements and one neuron of the same group showed a greater response during visual tracking than during fixation. Six out of 8 oriented cells with antagonistic areas showed almost the same response to the two types of visual stimulation, while the remaining two neurons showed very weak responses during smooth pursuit eye movements. Our results show that a small percentage (about 10%) of striate neurons in macaque monkeys gave very different responses to the same physical stimulation at retinal level, according to the presence or absence of slow eye movements (smooth pursuit eye movements). The activity of these neurons seems to be related to the real movement of something in the visual world, in spite of the retinal image movement per se.     

Binocular robot vision emulating disparity computation in the primary visual cortex

We designed a VLSI binocular vision system that emulates the disparity computation in the primary visual cortex (V1). The system consists of two silicon retinas, orientation chips, and field programmable gate array (FPGA), mimicking a hierarchical architecture of visual information processing in the disparity energy model. The silicon retinas emulate a Laplacian–Gaussian-like receptive field of the vertebrate retina. The orientation chips generate an orientation-selective receptive field by aggregating multiple pixels of the silicon retina, mimicking the Hubel–Wiesel-type feed-forward model in order to emulate a Gabor-like receptive field of simple cells. The FPGA receives outputs from the orientation chips corresponding to the left and right eyes and calculates the responses of the complex cells based on the disparity energy model. The system can provide the responses of complex cells tuned to five different disparities and a disparity map obtained by comparing these energy outputs. Owing to the combination of spatial filtering by analog parallel circuits and pixel-wise computation by hard-wired digital circuits, the present system can execute the disparity computation in real time using compact hardware.     

NR2B-subunit dependent facilitation of long-term potentiation in primary visual cortex following visual discrimination training of adult rats

Long‐term potentiation (LTP) is an important mechanism thought to mediate changes in synaptic connectivity following various types of experience. We examined the effects of visual discrimination training on LTP in the mature, rodent thalamocortical visual system. Adult rats underwent visual discrimination training in a modified Morris Water Maze containing a Y‐maze insert, requiring rats to associate visual cues with the location of a hidden escape platform placed in one of the two goal arms of the Y‐maze insert. On the day following successful task acquisition (average of nine training days), rats were anesthetized (urethane), and LTP in the thalamocortical system was characterized. In task‐naïve rats, theta‐burst stimulation of the lateral geniculate nucleus resulted in modest (～40%) potentiation of field postsynaptic potentials recorded in the primary visual cortex (V1). Rats trained on the visual discrimination task showed significantly greater levels of LTP (～60%), an effect that was not seen in rats trained to swim in the maze without a predictive association between visual cues and platform location. An antagonist of the N‐methyl‐d ‐aspartate (NMDA) receptor NR2B subunit ([R‐(R *,S *)]‐α‐(4‐hydroxyphenyl)‐β‐methyl‐4‐(phenylmethyl)‐1‐piperidinepropanol hydrochloride (Ro 25‐6981); 2 mm , applied locally at the recording site in V1) reversed the training‐induced LTP enhancement without affecting LTP in task‐naïve rats. An antagonist of metabotropic glutamate receptors [(2S)‐2‐amino‐2‐[(1S,2S)‐2‐carboxycycloprop‐1‐yl]‐3‐(xanth‐9‐yl) propanoic acid (LY 341495); 2 mm ] was ineffective in reversing the training‐induced LTP facilitation. These data suggest that behavioral (visual) training can result in changes in plasticity exhibited by the mature, thalamocortical visual system that require activation of NMDA receptors containing the NR2B subunit.     

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

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