Gratings Of Different Orientations Research Articles

Contrast sensitivity, visual acuity, and the effect of behavioural state on optokinetic gain in fiddler crabs.

Most animals rely on visual information for a variety of everyday tasks. The information available to a visual system depends in part on its spatial resolving power and contrast sensitivity. Due to their competing demands for physical space within an eye, these traits cannot simultaneously be improved without increasing overall eye size. The contrast sensitivity function is an integrated measure of visual performance that measures both resolution and contrast sensitivity. Its measurements helps us identify how different species have made a trade-off between contrast sensitivity and spatial resolution. It further allows us to identify the evolutionary drivers of sensory processing and visually mediated behaviour. Here, we measured the contrast sensitivity function of the fiddler crab Gelasimus dampieri using their optokinetic responses to wide-field, moving, sinusoidal intensity gratings of different orientations, spatial frequencies, contrasts, and speeds. We further tested whether the behavioural state of the crabs (i.e. whether crabs are actively walking or not) affects their optokinetic gain and contrast sensitivity. Our results from a group of five crabs suggest a minimum perceived contrast of 6%, and a horizontal and vertical visual acuity of 0.4 cyc/deg and 0.28 cyc/deg respectively in the crabs' region of maximum optomotor sensitivity. Optokinetic gain increased in moving crabs compared to restrained crabs, adding another example of the importance of naturalistic approaches when studying the performance of animals.

Effects of phase synchronization and frequency specificity in the encoding of conditioned fear-a web-based fear conditioning study.

Oscillatory synchronization in the theta-frequency band was found to play a causal role in binding information of different modalities in declarative memory. Moreover, there is first evidence from a laboratory study that theta-synchronized (vs. asynchronized) multimodal input in a classical fear conditioning paradigm resulted in better discrimination of a threat-associated stimulus when compared to perceptually similar stimuli never associated with the aversive unconditioned stimulus (US). Effects manifested in affective ratings and ratings of contingency knowledge. However, theta-specificity was not addressed so far. Thus, in the present pre-registered web-based fear conditioning study, we compared synchronized (vs. asynchronized) input in a theta-frequency band vs. the same synchronization manipulation in a delta frequency. Based on our previous laboratory design, five visual gratings of different orientations (25°, 35°, 45°, 55°, 65°) served as conditioned stimuli (CS) with only one (CS+) paired with the auditory aversive US. Both CS and US were luminance or amplitude modulated, respectively, in a theta (4 Hz) or delta (1.7 Hz) frequency. In both frequencies, CS-US pairings were presented either in-phase (0° phase lag) or out-of-phase (90°, 180°, 270°), resulting in four independent groups (each group N = 40). Phase synchronization augmented the discrimination of CSs in CS-US contingency knowledge but did not affect valence and arousal ratings. Interestingly, this effect occurred independent of frequency. In sum, the current study proves the ability to successfully conduct complex generalization fear conditioning in an online setting. Based on this prerequisite, our data supports a causal role of phase synchronization in the declarative CS-US associations for low frequencies rather than in the specific theta-frequency band.

Contrast sensitivity is formed by visual experience and task demands

The contrast sensitivity function (CSF) is a fundamental signature of the visual system and has been measured extensively in numerous species. It is defined by measurements of the visibility threshold for sinusoidal gratings at all spatial frequencies. Here, we measured the CSF in artificial visual systems, namely deep neural networks (DNNs), using the same contrast discrimination paradigm as used in human psychophysical experiments. During training, networks are exclusively exposed to natural images, and the task is to identify which one of two input images has higher contrast. The contrast discriminator networks learn a linear classifier over frozen features of pretrained DNNs. At the testing stage, we measured the network's CSF by presenting sinusoidal gratings of different orientations and spatial frequencies. Our results demonstrate that the pretrained DNNs show the band-limited, inverted U-shaped CSF, that is characteristic of the human CSF. The exact shape of the DNNs' CSF appears to be task-dependent. The human CSF is better captured by scene segmentation DNNs than image classification ones (tested under identical settings such as the architecture and computational complexity). When the network was trained to discriminate the contrast in natural images from scratch (i.e., with no previous training task), a relatively flat CSF emerged, dissimilar to the human CSF. The environment of pretrained DNNs proved influential as well. Those with a diet of natural images, similar to human experience (e.g. objects or faces), obtain more human-like CSF. Pretrained DNNs with aerial pictures, such as a bird flying in the sky would see, obtain a shifted CSF towards higher spatial frequencies, similar to that of an eagle. In conclusion, the CSFs derived from the DNNs are the result of efficient processing of the natural world around us. Different visual environments have succinct effects on the shape of the CSFs.

Pigeons (Columba livia) integrate visual motion using the vector average rule: effect of viewing distance.

Integrating local motion signals detected by the primary motion detector is crucial for representing a rigid, two-dimensional motion. The nature of motion integration has been studied using stimuli consisting of two superimposed sinusoidal gratings of different orientations, called plaid motion, and it has been shown that humans perceive integrated motion in the direction where the component constraint lines are intersected. We previously found that pigeons and humans perceive different movement directions from plaid motion; pigeons responded to the vector average direction of the gratings. Although this suggests that the underlying processes of motion integration differ between the two species, the viewing distance in the pigeon experiment, which used a touch panel procedure, was much smaller than in typical human experiments. The current study investigated the potential effect of viewing distance on the perception of plaid motion in pigeons. We trained six pigeons to detect whether motion directions were tilted leftward or rightward while a visual display was presented 0 or 40cm from an operant chamber. The pigeons responded to plaid stimuli for both viewing distance conditions as if they perceived motion in the vector average direction of two-component gratings. The result indicates that the species difference in plaid perception is not an artefact of viewing distance and suggests that pigeons use a different strategy than humans for integrating visual motion.

Plaid Detectors in Macaque V1 Revealed by Two-Photon Calcium Imaging.

Neuronal responses to one-dimensional orientations are combined to represent two-dimensional composite patterns; this plays a key role in intermediate-level vision such as texture segmentation. However, where and how the visual cortex starts to represent composite patterns, such as a plaid consisting of two superimposing gratings of different orientations, remains neurophysiologically elusive. Psychophysical and modeling evidence has suggested the existence of early neural mechanisms specialized in plaid detection [1-6], but the responses of V1 neurons to an optimally orientated grating are actually suppressed by a superimposing grating of different orientation (i.e., cross-orientation inhibition) [7, 8]. Would some other V1 neurons be plaid detectors? Here, we used two-photon calcium imaging [9] to compare the responses of V1 superficial-layer neurons to gratings and plaids in awake macaques. We found that many non-orientation-tuned neurons responded weakly to gratings but strongly to plaids, often with plaid orientation selectivity and cross-angle selectivity. In comparison, most (∼94%) orientation-tuned neurons showed more or less cross-orientation inhibition, regardless of the relative stimulus contrasts. Only a small portion (∼8%) of them showed plaid facilitation at off-peak orientations. These results suggest separate subpopulations of plaid and grating responding neurons. Because most of these plaid neurons (∼95%) were insensitive to motion direction, they were plaid pattern detectors, not plaid motion detectors.

Improving the signal-to-noise ratio of the visual P300

The P3b (“P300”) [1] of the event-related potential is considered to reflect cognitive —or even conscious [2]— stimulus processing. P300 recording sessions employ an “oddball paradigm”: target stimuli appear only infrequently among non-target stimuli. Thus they are quite lengthy and difficult to use with patients. We tested different strategies for minimizing the time needed to record a reliable visual P300. Targets were gratings of different orientation for target and non-target stimuli. Since previous studies only assessed absolute amplitude [e.g., 3, 4], we first determined the optimal signal-to-noise ratio for different target infrequencies. We found target-to-non-target ratios of 1:4 to 1:8 to be optimal. We next assessed whether shrinking the interval between stimulus onsets from 1000 ms to 214 ms would improve the signal-to-noise ratio when the recording time is kept constant, despite a temporal overlap of responses. This was indeed the case by a margin of up to 60%, but depended on the presentation duration of the stimuli. Finally, we assessed whether it is feasible to record a P300 in a manner similar to a steady-state visual evoked potential with a fixed number of intervening non-targets. We found a high inter-individual variability, but could reliably identify responses in all subjects with a multi-harmonic frequency-space analysis. Compared to conventional P300 recordings, less recording time was needed to acquire significant responses. In conclusion, the signal-to-noise ratio and the efficiency of visual P300 recordings can be improved by choosing optimal target infrequencies and by employing rapid stimulation schemes. [1] Linden, D. E. J. (2005). Neuroscientist 11:563-576. [2] Kotchoubey, B. (2005). Progr Brain Res 150:427-444. [3] Johnson, Jr. R. (1986). Psychophysiology 23:367-384. [4] Polich, J., & Margala, C. (1997). Int J Psychophysiol 25:169-176.

Quantification of Optical Images of Cortical Responses for Inferring Functional Maps

Optical imaging of cortical signals enables the mapping of functional organization across large patches of cortex with good spatial resolution. But techniques for the quantitative analysis and interpretation of these images are limited. Frequently the functional architecture of the cortex is inferred from the visible topography of cortical reflectance images averaged or differenced across stimulus conditions and scaled or color-coded for presentation. Such qualitative assessments have sometimes led to divergent conclusions particularly about the organization of spatial and temporal frequency preferences in the primary visual cortex. We applied quantitative methods derived from signal detection theory to objectively interpret optical images. The differential response to any two arbitrary stimuli was represented at each pixel as the probability of discriminating between the two stimuli given the reflectance values at that pixel. These probability maps reduced false alarms and provided better signal-to-noise ratio in fewer trials than difference maps. We applied these methods to optical images of primate primary visual area (V1) obtained in response to sinusoidal gratings of different orientations and spatiotemporal frequencies. Clustering by orientation preference was stronger than that for spatial frequency, whereas clustering by temporal frequency preference was the weakest, largely in agreement with a previous electrophysiological study that quantified the degree of clustering of neurons for various response properties using uniform, quantitative criterion. We suggest that probability maps can extend the applicability of optical imaging to investigations of finer aspects of cortical functional organization through better signal-to-noise ratio and uniform, quantitative criteria for interpretation.

Delayed discrimination of spatial frequency for gratings of different orientation: behavioral and fMRI evidence for low-level perceptual memory stores in early visual cortex

The concept of perceptual memory refers to the neural and cognitive processes underlying the storage of specific stimulus features such as spatial frequency, orientation, shape, contrast, and color. Psychophysical studies of perceptual memory indicate that observers can retain visual information about the spatial frequency of Gabor patterns independent of the orientation with which they are presented. Compared to discrimination of gratings with the same orientation, reaction times to orthogonally oriented gratings, however, increase suggesting additional processing. Using event-related fMRI we examined the pattern of neural activation evoked when subjects discriminated the spatial frequency of Gabors presented with the same or orthogonal orientation. Blood-oxygen level dependent BOLD fMRI revealed significantly elevated bilateral activity in visual areas (V1, V2) when the gratings to be compared had an orthogonal orientation, compared to when they had the same orientation. These findings suggest that a change in an irrelevant stimulus dimension requires additional processing in primary and secondary visual areas. The finding that the task-irrelevant stimulus property (orientation) had no significant effect on the prefrontal and intraparietal cortex supports a model of working memory in which discrimination and retention of basic stimulus dimensions is based on low-level perceptual memory stores that are located at an early stage in the visual process. Our findings suggest that accessing different stores requires time and has higher metabolic costs.

Orientation selectivity for sinusoidal gratings; evidence for an optical component

The human visual system exhibits lower contrast thresholds for detecting vertical and horizontal gratings compared with those orientated obliquely. This is known as the oblique effect and has been thought to be due wholly to neural factors. In this study, sensitivity to gratings of different orientation is compared when viewing with small and large pupils. Optical aberrations increase with pupil size. In some subjects, we find a statistically significant change in sensitivity to some orientations for the large pupil condition. This suggests that there is an optical component to orientation selectivity induced by the higher-order aberrations of the eye.

Directional responses of visual wulst neurones to grating and plaid patterns in the awake owl

The avian retinothalamofugal pathway reaches the telencephalon in an area known as visual wulst. A close functional analogy between this area and the early visual cortex of mammals has been established in owls. The goal of the present study was to assess quantitatively the directional selectivity and motion integration capability of visual wulst neurones, aspects that have not been previously investigated. We recorded extracellularly from a total of 101 cells in awake burrowing owls. From this sample, 88% of the units exhibited modulated directional responses to sinusoidal gratings, with a mean direction index of 0.74 +/- 0.03 and tuning bandwidth of 28 +/- 1.16 degrees . A direction index higher than 0.5 was observed in 66% of the cells, thereby qualifying them as direction selective. Motion integration was tested with moving plaids, made by adding two sinusoidal gratings of different orientations. We found that 80% of direction-selective cells responded optimally to the motion direction of the component gratings, whereas none responded to the global motion of plaids, whose direction was intermediate to that of the gratings. The remaining 20% were unclassifiable. The strength of component motion selectivity rapidly increased over a 200 ms period following stimulus onset, maintaining a relatively sustained profile thereafter. Overall, our data suggest that, as in the mammalian primary visual cortex, the visual wulst neurones of owls signal the local orientated features of a moving object. How and where these potentially ambiguous signals are integrated in the owl brain might be important for understanding the mechanisms underlying global motion perception.

'Simplification' of responses of complex cells in cat striate cortex: suppressive surrounds and 'feedback' inactivation.

In mammalian striate cortex (V1), two distinct functional classes of neurones, the so-called simple and complex cells, are routinely distinguished. They can be quantitatively differentiated from each other on the basis of the ratio between the phase-variant (F1) component and the mean firing rate (F0) of spike responses to luminance-modulated sinusoidal gratings (simple, F1/F0 > 1; complex, F1/F0 < 1). We investigated how recurrent cortico-cortical connections affect the spatial phase-variance of responses of V1 cells in the cat. F1/F0 ratios of the responses to optimally oriented drifting sine-wave gratings covering the classical receptive field (CRF) of single V1 cells were compared to those of: (1) responses to gratings covering the CRFs combined with gratings of different orientations presented to the 'silent' surrounds; and (2) responses to CRF stimulation during reversible inactivation of postero-temporal visual (PTV) cortex. For complex cells, the relative strength of the silent surround suppression on CRF-driven responses was positively correlated with the extent of increases in F1/F0 ratios. Inactivation of PTV cortex increased F1/F0 ratios of CRF-driven responses of complex cells only. Overall, activation of suppressive surrounds or inactivation of PTV 'converted' substantial proportions (50 and 30%, respectively) of complex cells into simple-like cells (F1/F0 > 1). Thus, the simple-complex distinction depends, at least partly, on information coming from the silent surrounds and/or feedback from 'higher-order' cortices. These results support the idea that simple and complex cells belong to the same basic cortical circuit and the spatial phase-variance of their responses depends on the relative strength of different synaptic inputs.

The Nature of V1 Neural Responses to 2D Moving Patterns Depends on Receptive-Field Structure in the Marmoset Monkey

A plaid pattern is formed when two sinusoidal gratings of different orientations are added together. Previous work has shown that V1 neurons selectively encode the direction and orientation of the component gratings in a moving plaid but not the direction of the plaid itself (Movshon et al. 1985). We recorded the responses of 49 direction-selective neurons to moving gratings and plaid patterns in area V1 of the anesthetized marmoset monkey (Callithrix jacchus). The responses of V1 neurons to rectangular patches of varying lengths and widths containing gratings of optimal spatial frequency were used to measure size and aspect ratio of the receptive-field subunits. We measured responses to plaid patterns moving in different directions and graded the magnitude of the response to the direction of motion of the plaid and the response to the direction of motion of the component gratings. We found significant correlations between receptive-field structure and the type and strength of its response to moving plaid patterns. The strength of pattern and component responses was significantly correlated with the interrelated properties of direction tuning width (Spearman's r = 0.82, P < 0.001), and receptive-field subunit aspect ratio (Spearman's r = -0.79, P < 0.001). Neurons with broad direction tuning and short, wide receptive-field subunits gave their greatest response when the plaid moved in their preferred direction. Conversely, neurons with narrow direction tuning and long, narrow receptive-field subunits gave their greatest responses when the plaid moved in a direction such that one of its components moved in the preferred direction.

A novel mechanism of response selectivity of neurons in cat visual cortex.

The spiking of cortical neurons critically depends on properties of the afferent stimuli. In the visual cortex, neurons respond selectively to the orientation and direction of movement of an object. The orientation and direction selectivity is improved upon transformation of the membrane potential changes into trains of action potentials. To address the question of whether the transformation of the membrane potential changes into spiking of a cell depends on the stimulus orientation and the direction of movement, we made intracellular recordings from the cat visual cortex in vivo during presentation of moving gratings of different orientations. We found that the relationship between the membrane polarization and the firing rate (input-output transfer function) depended on the stimulus orientation. The input-output transfer function was steepest during responses to the optimal stimulus; membrane depolarization of a given amplitude led to generation of more action potentials when evoked by an optimal stimulus than during non-optimal stimulation. The threshold for the action potential generation did not depend on stimulus orientation, and thus could not account for the observed difference in the transfer function. Oscillations of the membrane potential in the gamma-frequency range (25-70 Hz) were most pronounced during optimal stimulation and their strength changed in parallel with the changes in the transfer function, suggesting a possible relationship between the two parameters. We suggest that the improved input-output relationship of neurons during optimal stimulation represents a novel mechanism that may contribute to the final sharp orientation selectivity of spike responses in the cortical cells.

Learned discrimination of pattern orientation in walking flies.

To determine the pattern-orientation discrimination ability of blowflies, Phaenicia sericata, a learning/memory assay was developed in which sucrose served as the reward stimulus and was paired with one of two visual gratings of different orientations. Individual, freely walking flies with clipped wings were trained to discriminate between pairs of visual patterns presented in the vertical plane. During training trials, individual flies learned to search preferentially at the rewarded stimulus. In subsequent testing trials, flies continued to exhibit a learned preference for the previously rewarded stimulus, demonstrating an ability to discriminate between the two visual cues. Flies learned to discriminate between horizontal and vertical gratings, +45 degrees (relative to a 0 degrees vertical) and -45 degrees gratings, and vertical and +5 degrees gratings. Individual patterns of learning and locomotive behavior were observed in the pattern of exploration during training trials. The features of the visual cue critical for discrimination of orientation are discussed.

Receptive field properties of single neurons in rat primary visual cortex.

The rat is used widely to study various aspects of vision including developmental events and numerous pathologies, but surprisingly little is known about the functional properties of single neurons in the rat primary visual cortex (V1). These were investigated in the anesthetized (Hypnorm-Hypnovel), paralyzed animal by presenting gratings of different orientations, spatial and temporal frequencies, dimensions, and contrasts. Stimulus presentation and data collection were automated. Most neurons (190/205) showed sharply tuned (</=30 degrees bandwidth at half height) orientation selectivity with a bias for horizontal stimuli (31%). Analysis of response modulation of oriented cells showed a bimodal distribution consistent with the distinction between simple and complex cell types. Orientation specific interactions occurred between the center and the periphery of receptive fields, usually resulting in strong inhibition to center stimulation when both stimuli had the same orientation. There was no evidence for orientation columns nor for orderly change in optimal orientation with tangential tracks through V1. Responses were elicited by spatial frequencies ranging from zero (no grating) to 1.2 cycle/degree (c/ degrees ), peaking at 0.1 c/ degrees, and with a modal cutoff of 0.6 c/ degrees. Half of the neurons responded optimally to drifting gratings rather than flashing uniform field stimuli. Directional preference was seen for 59% of oriented units at all depths in the cortex. Optimal stimuli velocities varied from 10 to 250 degrees /s. Some units, mainly confined to layer 4, responded to velocities as high as 700 degrees /s. Response versus contrast curves (best fit with Naka-Rushton) varied from nearly linear to extremely steep (mean contrast semisaturation 50% and threshold 6%). There was a trend for cells from superficial layers to be more selective to different stimulus parameters than deeper layers cells. We conclude that neurons in rat V1 have complex and diverse visual properties, necessary for precise visual form perception with low spatial resolution.

Neuronal responses to plaids.

The majority of neurons in the visual cortex are orientation selective. When presented with a plaid, i.e. a stimulus generated by adding two gratings of different orientations, these neurons respond to the individual gratings making up the plaid. However, there are some pattern selective neurons in Area MT of the monkey visual cortex which respond in accordance with the combined plaid. The present study used computer simulation to investigate the response properties of simulated MT neurons to orthogonal plaids. The MT neurons were simulated by first multiplying the outputs of conventional orientation selective V1 neurons and then normalizing the product. It was discovered that pattern selective responses may emerge when the outputs from two orientation selective neurons, which differ in optimal orientation by more than about 50 degrees, are combined in this manner. This demonstrates that pattern selectivity may be the result of a very simple although nonlinear mechanism.

Perceived contrast following adaptation to gratings of different orientations

Using a contrast matching procedure, we measured the perceived contrast of vertical test gratings after adapting to other gratings of either vertical or horizontal orientation. The results show that both parallel and orthogonal adapting gratings reduce perceived contrast and do so proportionally more at low test contrasts than at high. The results are consistent with a single mechanism model proposed by Ross and Speed [(1991). Proceedings of the Royal Society (Series B), 246, 61-69] that assumes that adaptation to gratings repositions contrast-response transducer functions. They are not consistent with the notion of two different forms of adaptation, subtractive for parallel and multiplicative for orthogonal adaptors as proposed by Snowden and Hammett [(1992). Nature, 355, 248-250]. Nowhere is the reduction in perceived contrast by an orthogonal grating greater than that by a parallel grating of the same contrast. A direct comparison using two orthogonal adaptors confirms the greater potency of parallel adaptors, but also reveals interactions between the adaptors.

Interocular suppression in the primary visual cortex: a possible neural basis of binocular rivalry

In an attempt to demonstrate a physiological basis for the alternating suppression of perception when the two eyes view very different contours (binocular rivalry), we studied the responses of neurons in the lateral geniculate nucleus (LGN) and area 17 of 17 cats for drifting gratings of different orientation, spatial frequency and contrast in the two eyes. Almost half of the LGN neurons studied exhibited modest inhibitory interocular interaction, but independent of interocular differences in orientation. Monocularly driven units in layer 4 of area 17 behaved similarly. However, for the majority of binocular cortical cells, the response to a grating of optimal orientation in one eye was suppressed by a grating of very different orientation shown to the other eye, over a wide range of spatial frequency and independent of relative spatial phase. This interocular suppression exhibits a remarkable non-linearity: a grating of non-preferred orientation in one eye causes significant interocular suppression only if the neuron is already responding to an appropriate stimulus in the other eye [Sengpiel and Blakemore (1994) Nature, 368, 847–850]. We propose that the switches in perceptual dominance during binocular rivalry depend on interocular interactions at the level of binocular neurons of the primary visual cortex, which might involve intracortical inhibition between adjacent ocular dominance columns. The spontaneous alternations in perceptual suppression that occur during prolonged viewing of rivalrous patterns remain to be explained, although significant variation in the strength of neuronal suppression in such conditions was occasionally seen.

Interocular suppression in the visual cortex of strabismic cats

Strabismic humans usually experience powerful suppression of vision in the nonfixating eye. In an attempt to demonstrate physiological correlates of such suppression, we recorded from the primary visual cortex of cats with surgically induced squint and studied the responses of neurons to drifting gratings of different orientation, spatial frequency, and contrast in the two eyes. Only 1 of 50 apparently monocular cells showed any evidence of remaining, subliminal excitatory input from the "silent" eye when the two eyes were stimulated with gratings of similar orientation, and even among the small proportion of cells that remained binocularly driven, very few exhibited facilitation when stimulated binocularly. The majority of cells from both exotropes and esotropes, even those that could be independently driven through either eye, displayed nonspecific interocular suppression: stimulation of the nondominant eye with a drifting grating of any orientation depressed the response to an optimal grating being presented to the dominant eye. This phenomenon exhibited a gross nonlinearity in that it was dependent on the temporal sequence of stimulus presentation: stimulation of the nondominant eye caused significant suppression only if the neuron was already responding to an appropriate stimulus in the dominant eye, but not when onset of stimulation in the two eyes was simultaneous. Interocular suppression was always independent of the relative spatial phase of the two grating stimuli, and usually broadly tuned for the spatial frequency of the suppressive stimulus. Suppression may depend on inhibitory interaction between neighboring ocular dominance columns, combined with the loss of conventional disparity-selective binocular interactions for matched stimuli in the two eyes. The similarity of interocular suppression in strabismic cats and that caused by orthogonal gratings in the two eyes in normal cats (Sengpiel and Blakemore, 1994; Sengpiel et al., 1994) suggests that strabismic suppression and binocular rivalry depend on similar neural mechanisms.

The layout of iso-orientation domains in area 18 of cat visual cortex: optical imaging reveals a pinwheel-like organization

In this study we used optical imaging based on activity-dependent intrinsic signals to determine the distribution of cells responding to gratings of various orientations moving in different directions in area 18 of cat visual cortex. To test directional-selective clustering of neurons, we compared cortical activity maps obtained by stimulation with two gratings of identical orientation but moving in opposite directions. We found those maps to be almost identical, suggesting that neurons are not notably clustered into directionality columns. We also compared activity maps obtained with gratings of different orientations. Each of the orientation maps was similar to the 2-deoxyglucose maps previously reported. Having compiled the information obtained from the different orientations into one "orientation preference map," we found, in contrast to earlier reports, that iso-orientation domains are not elongated parallel bands but are small patches organized in "pinwheels" around points that we refer to as "orientation centers." We furthermore show that the only locations at which orientation preference changes rapidly are these orientation centers and not lines or loops. In addition, this report clarifies that our observations on the functional architecture of cat area 18, although at first sight at variance with earlier observations, are actually fully consistent with them. We therefore propose that in cat visual cortex pinwheel-like patterns of orientation preference form an irregular mosaic of modular units with an average density of 1.2 pinwheels per square millimeter.