Network‐selectivity and stimulus‐discrimination in the primary visual cortex: cell‐assembly dynamics

Visual neurons coordinate their responses in relation to the stimulus; however, the complex interplay between a stimulus and the functional dynamics of an assembly still eludes neuroscientists. To this aim, we recorded cell assemblies from multi‐electrodes in the primary visual cortex of anaesthetized cats in response to randomly presented sine‐wave drifting gratings whose orientation tilted in 22.5° steps. Cross‐correlograms revealed the functional connections at all the tested orientations. We show that a cell‐assembly discriminates between orientations by recruiting a ‘salient’ functional network at every presented orientation, wherein the connections and their strengths (peak‐probabilities in the cross‐correlogram) change from one orientation to another. Within these assemblies, closely tuned neurons exhibited increased connectivity and connection‐strengths compared with differently tuned neurons. Minimal connectivity between untuned neurons suggests the significance of neuronal selectivity in assemblies. This study reflects upon the dynamics of functional connectivity, and brings to the fore the importance of a ‘signature’ functional network in an assembly that is strictly related to a specific stimulus. It appears that an assembly is the major ‘functional unit’ of information processing in cortical circuits, rather than the individual neurons.     

The effect of task‐irrelevant visual backgrounds on human transcranial magnetic stimulation‐evoked electroencephalography responses and cortical alpha activity

Brain responses evoked by transcranial magnetic stimulation (TMS) in task‐free experimental contexts are known to depend on psychophysiological states such as sleep, vegetative state and caffeine‐induced arousal. Much less is known about how TMS‐evoked responses depend on task‐irrelevant steady perceptual input. Here, we examined ongoing alpha activity and the mean amplitude of EEG potentials in response to occipitally applied TMS as a function of task‐irrelevant visual backgrounds. Responses to TMS were robustly modulated by photographs of natural scenes and man‐made environments. These effects began as early as during the N100 and continued for several hundred milliseconds after the stimulation. There was also a more general effect of background along with other stimuli, such as blank backgrounds, sinusoidal gratings and moving dot‐patterns. This effect was observable from ongoing alpha activity as well. Based on these results we conclude that different types of steady perceptual input modulate visual cortex reactivity and/or connectivity and it is possible to measure these modulations by combining TMS with electroencephalography.     

Human area V5 and motion in the ipsilateral visual field

We have studied area V5 of the human brain with visually-evoked potential (VEP) and functional magnetic resonance imaging (fMRI) methods, using hemifield motion stimuli. Our results confirmed the presence of an ipsilateral field representation in V5 and found: (i) a delay in the ipsilateral response in V5, irrespective of the hemifield stimulated; (ii) a longer ipsilateral delay for left hemifield than for right hemifield stimulation; and (iii) in a patient with a section of the splenium, an absent ipsilateral response for right but not left hemifield stimulation. Together with neurophysiological and anatomical evidence in the monkey, our non-invasive spatial and temporal imaging studies in man reveal that ipsilateral V5 is activated by motion signals transferred from contralateral V5. The asymmetry of ipsilateral delay in normal subjects and the asymmetrical loss of ipsilateral response following splenial section imply that signals related to visual motion are transferred from one V5 to the other through two segregated pathways.     

Differential brain mechanisms for processing distracting information in task‐relevant and ‐irrelevant dimensions in visual search

A crucial function of our goal‐directed behavior is to select task‐relevant targets among distractor stimuli, some of which may share properties with the target and thus compete for attentional selection. Here, by applying functional magnetic resonance imaging (fMRI) to a visual search task in which a target was embedded in an array of distractors that were homogeneous or heterogeneous along the task‐relevant (orientation or form) and/or task‐irrelevant (color) dimensions, we demonstrate that for both (orientation) feature search and (form) conjunction search, the fusiform gyrus is involved in processing the task‐irrelevant color information, while the bilateral frontal eye fields (FEF), the cortex along the left intraparietal sulcus (IPS), and the left junction of intraparietal and transverse occipital sulci (IPTO) are involved in processing task‐relevant distracting information, especially for target‐absent trials. Moreover, in conjunction (but not in feature) search, activity in these frontoparietal regions is affected by stimulus heterogeneity along the task‐irrelevant dimension: heterogeneity of the task‐irrelevant information increases the activity in these regions only when the task‐relevant information is homogeneous, not when it is heterogeneous. These findings suggest that differential neural mechanisms are involved in processing task‐relevant and task‐irrelevant dimensions of the searched‐for objects. In addition, they show that the top‐down task set plays a dominant role in determining whether or not task‐irrelevant information can affect the processing of the task‐relevant dimension in the frontoparietal regions.     

Direction selectivity of neurons in the visual cortex is non‐linear and lamina‐dependent

Neurons in the visual cortex are generally selective to direction of movement of a stimulus. Although models of this direction selectivity (DS) assume linearity, experimental data show stronger degrees of DS than those predicted by linear models. Our current study was intended to determine the degree of non‐linearity of the DS mechanism for cells within different laminae of the cat's primary visual cortex. To do this, we analysed cells in our database by using neurophysiological and histological approaches to quantify non‐linear components of DS in four principal cortical laminae (layers 2/3, 4, 5, and 6). We used a DS index (DSI) to quantify degrees of DS in our sample. Our results showed laminar differences. In layer 4, the main thalamic input region, most neurons were of the simple type and showed high DSI values. For complex cells in layer 4, there was a broad distribution of DSI values. Similar features were observed in layer 2/3, but complex cells were dominant. In deeper layers (5 and 6), DSI value distributions were characterized by clear peaks at high values. Independently of specific lamina, high DSI values were accompanied by narrow orientation tuning widths. Differences in orientation tuning for non‐preferred vs. preferred directions were smallest in layer 4 and largest in layer 6. These results are consistent with a non‐linear process of intra‐cortical inhibition that enhances DS by selective suppression of neuronal firing for non‐preferred directions of stimulus motion in a lamina‐dependent manner. Other potential mechanisms are also considered.     

Segregation of visual inputs from different regions of the compound eye in two parallel pathways through the anterior optic tubercle of the bumblebee (Bombus ignitus)

Visually guided behaviors require the brain to extract features of the visual world and to integrate them in a context-specific manner. Hymenopteran insects have been prime models for ethological research into visual behaviors for decades but knowledge about the underlying central processing is very limited. This is particularly the case for sky-compass navigation. To learn more about central processing of visual information in general and specifically to reveal a possible polarization vision pathway in the bee brain, we used tracer injections to investigate the pathways through the anterior optic tubercle, a prominent output target of the insect optic lobe, in the bumblebee Bombus ignitus. The anterior optic tubercle of the bumblebee is a small neuropil of 200 μm width and is located dorsolateral to the antennal lobe at the anterior surface of the brain. It is divided into a larger upper and a smaller lower subunit, both of which receive input from the optic lobe and connect to the lateral accessory lobe, and the contralateral tubercle, via two parallel pathways. The lower subunit receives input from the dorsal rim area (DRA) of the compound eye. The bumblebee DRA shares structural similarities with polarization-sensitive DRAs of other insects and looks similar to that of honeybees. We identified several neurons within this pathway that could be homologous to identified polarization-sensitive neurons in the locust brain. We therefore conclude that the pathway through the lower subunit of the anterior optic tubercle could carry polarization information from the periphery to the central brain.     

On the domain‐specificity of the visual and non‐visual face‐selective regions

What happens in our brains when we see a face? The neural mechanisms of face processing – namely, the face‐selective regions – have been extensively explored. Research has traditionally focused on visual cortex face‐regions; more recently, the role of face‐regions outside the visual cortex (i.e., non‐visual‐cortex face‐regions) has been acknowledged as well. The major quest today is to reveal the functional role of each this region in face processing. To make progress in this direction, it is essential to understand the extent to which the face‐regions, and particularly the non‐visual‐cortex face‐regions, process only faces (i.e., face‐specific, domain‐specific processing) or rather are involved in a more domain‐general cognitive processing. In the current functional MRI study, we systematically examined the activity of the whole face‐network during face‐unrelated reading task (i.e., written meaningful sentences with content unrelated to faces/people and non‐words). We found that the non‐visual‐cortex (i.e., right lateral prefrontal cortex and posterior superior temporal sulcus), but not the visual cortex face‐regions, responded significantly stronger to sentences than to non‐words. In general, some degree of sentence selectivity was found in all non‐visual‐cortex cortex. Present result highlights the possibility that the processing in the non‐visual‐cortex face‐selective regions might not be exclusively face‐specific, but rather more or even fully domain‐general. In this paper, we illustrate how the knowledge about domain‐general processing in face‐regions can help to advance our general understanding of face processing mechanisms. Our results therefore suggest that the problem of face processing should be approached in the broader scope of cognition in general.     

The microglial fractalkine receptor is not required for activity‐dependent plasticity in the mouse visual system

Microglia have recently been implicated as key regulators of activity‐dependent plasticity, where they contribute to the removal of inappropriate or excess synapses. However, the molecular mechanisms that mediate this microglial function are still not well understood. Although multiple studies have implicated fractalkine signaling as a mediator of microglia–neuron communications during synaptic plasticity, it is unclear whether this is a universal signaling mechanism or whether its role is limited to specific brain regions and stages of the lifespan. Here, we examined whether fractalkine signaling mediates microglial contributions to activity‐dependent plasticity in the developing and adolescent visual system. Using genetic ablation of fractalkine's cognate receptor, CX₃CR1, and both ex vivo characterization and in vivo imaging in mice, we examined whether fractalkine signaling is required for microglial dynamics and modulation of synapses, as well as activity‐dependent plasticity in the visual system. We did not find a role for fractalkine signaling in mediating microglial properties during visual plasticity. Ablation of CX₃CR1 had no effect on microglial density, distribution, morphology, or motility, in either adolescent or young adult mice across brain regions that include the visual cortex. Ablation of CX₃CR1 also had no effect on baseline synaptic turnover or contact dynamics between microglia and neurons. Finally, we found that fractalkine signaling is not required for either early or late forms of activity‐dependent visual system plasticity. These findings suggest that fractalkine is not a universal regulator of synaptic plasticity, but rather has heterogeneous roles in specific brain regions and life stages.     

Stimulus‐dependent augmented gamma oscillatory activity between the functionally connected cortical neurons in the primary visual cortex

Neuronal assemblies typically synchronise within the gamma oscillatory band (30–80 Hz) and are fundamental to information processing. Despite numerous investigations, the exact mechanisms and origins of gamma oscillations are yet to be known. Here, through multiunit recordings in the primary visual cortex of cats, we show that the strength of gamma power (20–40 and 60–80 Hz) is significantly stronger between the functionally connected units than between the unconnected units within an assembly. Furthermore, there is increased frequency coherence in the gamma band between the connected units than between the unconnected units. Finally, the higher gamma rhythms (60–80 Hz) are mostly linked to the fast‐spiking neurons. These results led us to postulate that gamma oscillations are intrinsically generated between the connected units within cell assemblies (microcircuits) in relation to the stimulus within an emergent ‘50‐ms temporal window of opportunity’.     

Distribution of calcium-binding proteins within the parallel visual pathways of a primate (Galago crassicaudatus)

Bush babies possess three distinct parallel pathways to striate cortex (V1 or area 17). The calcium-binding proteins parvalbumin (PV) and calbindin (CB) typically show complementary regional distributions in the brain, often associated with specific aspects of functionally related groups of cells. We asked whether PV⁺ and CB⁺ immunoreactivity differentiate central visual parallel pathways in this species. Results show that PV and CB cell and neuropil staining is strongly complementary in the lateral geniculate nucleus (LGN) and is associated with separate parallel pathways. CB⁺ immunoreactivity is dense, but cytochrome oxidase (CO) staining is light in the paired koniocellular layers. PV⁺ and CO⁺ immunoreactivity is most dense in the parvocellular and magnocellular layers. Combined analyses of cell size, retrograde labeling, and double labeling have confirmed that all PV⁺ and CB⁺ LGN cells are geniculocortical relay cells; none was found to be σ-aminobutyric acid (GABA)ergic. In V1, dense PV⁺ neuropil closely matches the expression of CO in layer 4 and in the blobs of layer 3. CB⁺ staining is most dense in layers 2 and 3A and is not strongly expressed within the CO interblobs. Finally, PV and CB are not found in related parallel pathway components in the LGN and V1 (e. g., in V1, CO blobs exhibit dense PV⁺ neuropil, yet they are targets of the small K geniculocortical relay cells that are CB⁺ in the LGN). Our findings support the view that three functionally distinct visual pathways project to V1 from the LGN. However, the differences in the patterns of localization of PV and CB in the LGN and in V1 suggest that these proteins may be utilized in different ways in these two visual areas. © 1995 Wiley-Liss, Inc.     

LSD alters eyes‐closed functional connectivity within the early visual cortex in a retinotopic fashion

The question of how spatially organized activity in the visual cortex behaves during eyes‐closed, lysergic acid diethylamide (LSD)‐induced “psychedelic imagery” (e.g., visions of geometric patterns and more complex phenomena) has never been empirically addressed, although it has been proposed that under psychedelics, with eyes‐closed, the brain may function “as if” there is visual input when there is none. In this work, resting‐state functional connectivity (RSFC) data was analyzed from 10 healthy subjects under the influence of LSD and, separately, placebo. It was suspected that eyes‐closed psychedelic imagery might involve transient local retinotopic activation, of the sort typically associated with visual stimulation. To test this, it was hypothesized that, under LSD, patches of the visual cortex with congruent retinotopic representations would show greater RSFC than incongruent patches. Using a retinotopic localizer performed during a nondrug baseline condition, nonadjacent patches of V1 and V3 that represent the vertical or the horizontal meridians of the visual field were identified. Subsequently, RSFC between V1 and V3 was measured with respect to these a priori identified patches. Consistent with our prior hypothesis, the difference between RSFC of patches with congruent retinotopic specificity (horizontal–horizontal and vertical–vertical) and those with incongruent specificity (horizontal–vertical and vertical–horizontal) increased significantly under LSD relative to placebo, suggesting that activity within the visual cortex becomes more dependent on its intrinsic retinotopic organization in the drug condition. This result may indicate that under LSD, with eyes‐closed, the early visual system behaves as if it were seeing spatially localized visual inputs. Hum Brain Mapp 37:3031–3040, 2016. © 2016 Wiley Periodicals, Inc .     

Neural correlates of atomoxetine improving inhibitory control and visual processing in Drug‐naïve adults with attention‐deficit/hyperactivity disorder

Atomoxetine improves inhibitory control and visual processing in healthy volunteers and adults with attention‐deficit/hyperactivity disorder (ADHD). However, little is known about the neural correlates of these two functions after chronic treatment with atomoxetine. This study aimed to use the counting Stroop task with functional magnetic resonance imaging (fMRI) and the Cambridge Neuropsychological Test Automated Battery (CANTAB) to investigate the changes related to inhibitory control and visual processing in adults with ADHD. This study is an 8‐week, placebo‐controlled, double‐blind, randomized clinical trial of atomoxetine in 24 drug‐naïve adults with ADHD. We investigated the changes of treatment with atomoxetine compared to placebo‐treated counterparts using the counting Stroop fMRI and two CANTAB tests: rapid visual information processing (RVP) for inhibitory control and delayed matching to sample (DMS) for visual processing. Atomoxetine decreased activations in the right inferior frontal gyrus and anterior cingulate cortex, which were correlated with the improvement in inhibitory control assessed by the RVP. Also, atomoxetine increased activation in the left precuneus, which was correlated with the improvement in the mean latency of correct responses assessed by the DMS. Moreover, anterior cingulate activation in the pre‐treatment was able to predict the improvements of clinical symptoms. Treatment with atomoxetine may improve inhibitory control to suppress interference and may enhance the visual processing to process numbers. In addition, the anterior cingulate cortex might play an important role as a biological marker for the treatment effectiveness of atomoxetine. Hum Brain Mapp 38:4850–4864, 2017. © 2017 Wiley Periodicals, Inc.     

High‐level visual manifestations of epileptic seizures originating from the medial parietal cortex

We describe the case of a patient with well‐localized focal seizures originating from the medial parietal cortex. Seizures originated from area 7m, and findings revealed clear visuospatial semiological signs that may be used clinically to help diagnose similar cases of seizures in non‐lesional patients.     

Optic nerve,superior colliculus,visual thalamus,and primary visual cortex of the northern elephant seal (Mirounga angustirostris) and California sea lion (Zalophus californianus)

The northern elephant seal (Mirounga angustirostris) and California sea lion (Zalophus californianus) are members of a diverse clade of carnivorous mammals known as pinnipeds. Pinnipeds are notable for their large, ape‐sized brains, yet little is known about their central nervous system. Both the northern elephant seal and California sea lion spend most of their lives at sea, but each also spends time on land to breed and give birth. These unique coastal niches may be reflected in specific evolutionary adaptations to their sensory systems. Here, we report on components of the visual pathway in these two species. We found evidence for two classes of myelinated fibers within the pinniped optic nerve, those with thick myelin sheaths (elephant seal: 9%, sea lion: 7%) and thin myelin sheaths (elephant seal: 91%, sea lion: 93%). In order to investigate the architecture of the lateral geniculate nucleus, superior colliculus, and primary visual cortex, we processed brain sections from seal and sea lion pups for Nissl substance, cytochrome oxidase, and vesicular glutamate transporters. As in other carnivores, the dorsal lateral geniculate nucleus consisted of three main layers, A, A1, and C, while each superior colliculus similarly consisted of seven distinct layers. The sea lion visual cortex is located at the posterior side of cortex between the upper and lower banks of the postlateral sulcus, while the elephant seal visual cortex extends far more anteriorly along the dorsal surface and medial wall. These results are relevant to comparative studies related to the evolution of large brains.     

Topographic arrangement of the rotundo‐entopallial projection in the pigeon (Columba livia)

The tectofugal pathway (retina – optic tectum – nucleus rotundus – entopallium) is a prominent route mediating visual discrimination in diurnal birds. Several lines of evidence have shown that at the tecto‐rotundal stage this pathway is composed of multiple parallel channels. Anatomical studies show that the nucleus rotundus is composed of at least four subdivisions, according to differences in cytoarchitectonic, histochemical, and hodological properties. Each of these subdivisions is in receipt of a highly convergent, nontopographic tectal projection, originating from a distinct subset of tecto‐rotundal neurons. Physiological studies show that neurons of different subdivisions respond specifically to different visual dimensions, such as color, luminance, two‐dimensional motion, and in‐depth motion. At present it is less clear whether or to what extent this channel segregation is preserved at the telencephalic stage of the tectofugal pathway. The entopallium shows no obvious subdivisions or laminations. Nevertheless, tract‐tracing experiments show that separate portions of the entopallium receive efferent projections from different rotundal subdivisions, in a way that maintains the rostrocaudal order of these subdivisions. In the present study we investigate in detail the topography of the rotundo‐entopallial projection by means of anterograde and retrograde neuronal tracers. Our results confirm the zonal topography proposed by previous studies and indicate that each zone in the entopallium receives a direct and topographically organized projection from its corresponding rotundal subdivision. These results suggest that the spatial arrangement of the different rotundal functional modules is preserved at the entopallial level. J. Comp. Neurol. 518:4342–4361, 2010. © 2010 Wiley‐Liss, Inc.     

Top‐down and bottom‐up influences on the left ventral occipito‐temporal cortex during visual word recognition: An analysis of effective connectivity

The functional role of the left ventral occipito‐temporal cortex (vOT) in visual word processing has been studied extensively. A prominent observation is higher activation for unfamiliar but pronounceable letter strings compared to regular words in this region. Some functional accounts have interpreted this finding as driven by top‐down influences (e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254–262; Price and Devlin [ 2011 ]: Trends Cogn Sci 15:246–253), while others have suggested a difference in bottom‐up processing (e.g., Glezer et al. [ 2009 ]: Neuron 62:199–204; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594). We used dynamic causal modeling for fMRI data to test bottom‐up and top‐down influences on the left vOT during visual processing of regular words and unfamiliar letter strings. Regular words (e.g., taxi) and unfamiliar letter strings of pseudohomophones (e.g., taksi) were presented in the context of a phonological lexical decision task (i.e., “Does the item sound like a word?”). We found no differences in top‐down signaling, but a strong increase in bottom‐up signaling from the occipital cortex to the left vOT for pseudohomophones compared to words. This finding can be linked to functional accounts which assume that the left vOT contains neurons tuned to complex orthographic features such as morphemes or words [e.g., Dehaene and Cohen [ 2011 ]: Trends Cogn Sci 15:254‐262; Kronbichler et al. [ 2007 ]: J Cogn Neurosci 19:1584–1594]: For words, bottom‐up signals converge onto a matching orthographic representation in the left vOT. For pseudohomophones, the propagated signals do not converge, but (partially) activate multiple orthographic word representations, reflected in increased effective connectivity. Hum Brain Mapp 35:1668–1680, 2014. © 2013 Wiley Periodicals, Inc.     

Chemoarchitecture of the middle temporal visual area in the marmoset monkey (Callithrix jacchus): laminar distribution of calcium-binding proteins (calbindin, parvalbumin) and nonphosphorylated neurofilament

We studied the distributions of interneurons containing the calcium-binding proteins parvalbumin and calbindin D-28k, as well as that of pyramidal neurons containing nonphosphorylated neurofilament (NNF), in the middle temporal visual area (MT) of marmoset monkeys. The distributions of these classes of cells in MT are distinct from those found in adjacent areas. Similar to the primary visual area (V1), in MT, calbindin-immunopositive neurons can be objectively classified into "dark" and "light" subtypes based on optical density of stained cell bodies. Calbindin-positive dark neurons are particularly concentrated in layers 2 and 3, whereas light neurons have a more widespread distribution. In addition, a subcategory of calbindin-positive dark neuron, characterized by a "halo" of stained processes surrounding the cell body, is found within and around layer 4 of MT and V1. These cells are rare in most other visual areas. In comparison, parvalbumin-immunopositive cells in area MT have a relatively homogeneous distribution, although with a trend toward higher spatial density in lower layer 3, and are relatively uniform in terms of density of staining. Finally, MT shows a characteristic trilaminar distribution of NNF-immunopositive pyramidal cells, with stained cell bodies evident in layers 3, 5, and 6. Although the laminar distribution of cells stained for the three markers overlap to some extent, these subcategories can be readily distinguished in terms of morphology, including cell body size. Chemoarchitectural parallels observed between MT and V1 suggest comparable physiological requirements and neuronal circuitry.     

Representation of the visual field in the primary visual area of the marmoset monkey: Magnification factors,point‐image size,and proportionality to retinal ganglion cell density

The primary visual area (V1) forms a systematic map of the visual field, in which adjacent cell clusters represent adjacent points of visual space. A precise quantification of this map is key to understanding the anatomical relationships between neurons located in different stations of the visual pathway, as well as the neural bases of visual performance in different regions of the visual field. We used computational methods to quantify the visual topography of V1 in the marmoset (Callithrix jacchus), a small diurnal monkey. The receptive fields of neurons throughout V1 were mapped in two anesthetized animals using electrophysiological recordings. Following histological reconstruction, precise 3D reconstructions of the V1 surface and recording sites were generated. We found that the areal magnification factor (M_A) decreases with eccentricity following a function that has the same slope as that observed in larger diurnal primates, including macaque, squirrel, and capuchin monkeys, and humans. However, there was no systematic relationship between M_A and polar angle. Despite individual variation in the shape of V1, the relationship between M_A and eccentricity was preserved across cases. Comparison between V1 and the retinal ganglion cell density demonstrated preferential magnification of central space in the cortex. The size of the cortical compartment activated by a punctiform stimulus decreased from the foveal representation towards the peripheral representation. Nonetheless, the relationship between the receptive field sizes of V1 cells and the density of ganglion cells suggested that each V1 cell receives information from a similar number of retinal neurons, throughout the visual field. J. Comp. Neurol. 521:1001–1019, 2013. © 2012 Wiley Periodicals, Inc.     

Spatial-frequency tuning and geniculocortical projections in the visual cortex (areas 17 and 18) of the pigmented ferret

We have examined the spatial-frequency selectivity of neurons in areas 17 and 18 of the adult pigmented ferret, by measuring how the amplitude of response depends on the spatial-frequency of moving sinusoidal gratings of optimal orientation and fixed contrast. Neurons in area 17 of the ferret respond optimally to low spatial frequencies [average 0.25 cycles per degree (c/deg)], much lower than the optima for cat area 17. The tuning curves are of the same form as those found in cat and monkey: unimodal with bandwidths in the range 0.8–3.5 octaves. Neurons in area 18 of the ferret respond optimally to even lower spatial frequencies (average 0.087 c/deg) than area 17 neurons, and the distributions of optimal spatial frequency for areas 17 and 18 hardly overlap. In both cortical areas, the bandwidth of the tuning curves is inversely correlated with optimal spatial frequency. This marked difference in tuning between the two cortical areas is probably attributable to differential geniculo-cortical projections. Small injections of fluorescent latex microspheres or horseradish peroxidase (HRP) were made into area 17 or area 18 in order to investigate the populations of geniculate neurons projecting to the two cortical areas. After injections into area 17, labelled neurons are found predominantly in the geniculate A layers, with a few neurons labelled in the C layers. Conversely, after an area 18 injection, similar numbers of labelled neurons are found in the C layers as in the A layers. Soma-size analysis of the neurons in the A-layers suggests the existence of two populations of relay neurons, which project differentially to areas 17 and 18. The different geniculate inputs and the different spatial-frequency tuning in areas 17 and 18 may imply that the two cortical areas process visual information more in parallel than in series.