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Anisotropic local contrast normalization: the role of stimulus orientation and spatial frequency bandwidths in the oblique and horizontal effect perceptual anisotropies.

Visual ability for sine waves and other narrowband stimuli shows an oblique effect--worst performance at obliques, best at horizontal and vertical orientations. Recently, we have shown that with broadband stimuli (either 1/f(alpha) visual noise or natural scenes), performance for detecting oriented content is worst at horizontal, best at the obliques, and intermediate at vertical orientations (a "horizontal effect"). This horizontal effect has been explained by a cortical contrast normalization model that is both local (over orientation and spatial frequency) and anisotropic (due to a numerical bias of neurons with different preferred orientations). Here, the bandwidth of content at which an oblique effect or horizontal effect occurs was assessed in two suprathreshold matching experiments conducted with 1/f(alpha) noise stimuli filtered with a triangle increment function of varied bandwidth (16 levels of orientation and spatial frequency bandwidth). The results provided further support for the local anisotropic normalization model in that an oblique effect was observed when a fairly small range of orientations and high spatial frequencies were tested and the horizontal effect was observed for broadband increments > or = 20 degrees orientation bandwidth and > or = 1-octave in frequency. At intermediate spatial frequency and orientation increment bandwidths, a blend of the two anisotropies was observed.

Adult↗

Effects of spatial attention on detection and identification of oriented lines.

In this paper, we study the effects of spatial attention on detection and identification of oriented lines presented at near-threshold luminance. In the first experiment, we found that the cuing effect was greater when observers had to discriminate between two close orientations than when they had to discriminate between two far-apart orientations. In the second experiment, we examined the effects of peripheral cues on summation of low-contrast oriented lines. We found that the range of orientations within which summation occurred was greater when the cues were invalid than when they were valid. Our results suggest that attention to a specific location in the visual field modulates the neural channels which code orientation in at least two different ways: (a) spatial attention increases the responsivity of these channels to stimuli presented at this location, and (b) it reduces the bandwidths or ranges of orientations to which these channels are sensitive. These results suggest that the properties of the orientation-tuned channels, including those that seem to exert their effects at early stages of orientation processing (e.g., tuning function), may not be fixed, but rather vary according to the attention being paid to the spatial region within which the target stimulus is presented.

Adult↗

Attentional demands of continuously monitoring orientation using vestibular information.

The aim of this series of experiments was to determine whether attention is normally required for continuously processing vestibular information concerning orientation, or is required only when orientation is disrupted (eg by vestibular dysfunction or by conflicting visual and vestibular orientation cues). In the first two studies, healthy subjects were passively oscillated, and indicated when they perceived they were passing through their starting position. There was only weak evidence for interference between performance on this 'continuous orientation monitoring task' and on concurrent mental tasks. However, a third study showed that when patients with vestibular imbalance carried out the continuous orientation monitoring task their performance on a concurrent mental arithmetic task was substantially impaired. This dual task interference was correlated with inaccuracy in judging orientation on the continuous orientation monitoring task, which in turn correlated with severity of recent vestibular symptomatology (assessed by questionnaire). In a fourth experiment, disorientation was induced in healthy subjects by rotating the visual field about the line of sight. Bidirectional interference was observed between monitoring orientation (assessed by accuracy in setting a rod to the perceived vertical) and performance of an arithmetic task. Dual task interference was correlated with baseline levels of disorientation induced by the visual field, as indicated by inaccuracy in judging the visual vertical. These findings suggest that monitoring orientation makes significant demands upon cortical processing resources when disorientation is induced, whether the disorientation results from deficient sensory functioning or from ambiguous perceptual information.

Adult↗

Discrimination of an orientation difference in dynamic textures.

We investigated whether the response of a motion sensor was related to the specificity of sensory information (orientation and direction of motion) used to compute motion energy. This was done in two ways. First, we assessed whether orientation discrimination of a target line, which segregated by an orientation difference from a textured background, was improved with two-frame apparent motion stimulation (as compared with static presentation). Second, we investigated whether the amount of improvement (in either orientation or direction of motion discrimination) depends on a particular combination of target orientation and direction of motion (either orthogonal or parallel). We found that the percentage of correct responses in the discrimination task (a) was higher for a moving target than for a static one; (b) was higher when the target was oriented more orthogonally to motion direction than background elements; (c) was little affected by background motion and (d) decreased with frame duration in the direction of motion task whereas it was largely unaffected by frame duration in the discrimination of orientation task. These results suggest that discrimination of moving texture boundaries is based on a motion sensor tuned to a particular combination of orientation and direction of motion, which is capable of signalling the orientation of a moving target more accurately than a static sensor.

Analysis of Variance↗

Orientation sensitivity in human visual motion processing.

Orientation tuning of receptive fields is well documented in the spatial domain, but considerable variability exists amongst published estimates of orientation sensitivity of motion receptive fields. We used a two-frame motion sequence, in which one frame was binary noise and the other was a horizontally displaced and filtered version of the same noise field, to examine the orientation sensitivity of human motion mechanisms. Initially, orientations orthogonal to the direction of motion were removed from each filtered frame. Observers indicated perceived direction of motion in a single interval, binary choice task. D(max) was determined for different amounts of removed orientations, and found to remain constant across the removal of energy up to approximately +/-60 deg from vertical. In a second experiment, the orientations removed were now parallel to the direction of motion of the stimulus. D(max) fell as a cosine function with increasing removal of orientation information, in agreement with off-orientation looking or matched filtering predictions. The two experiments show the presence of mechanisms both broadly tuned and more narrowly tuned for orientation. A control experiment introduced an interstimulus interval between the two frames of our motion sequence. Performance on the direction discrimination task was severely degraded, indicating that the original results are not explicable in terms of a feature-tracking or long-range motion process. The presence of both broadly and narrowly tuned mechanisms implies multiple possible solutions to the processing of coherent plaid motion.

Humans↗

Brainstem control of head movements during orienting; organization of the premotor circuits.

When an object appears in the visual field, animals orient their head, eyes, and body toward it in a well-coordinated manner (orienting movement). The head movement is a major portion of the orienting movement. Interest in the neural control of head movements in the monkey and human have increased in the 1990's, however, fundamental knowledge about the neural circuits controlling the orienting head movement continues to be based on a large number of experimental studies performed in the cat. Thus, it is crucial now to summarize information that has been clarified in the cat for further advancement in understanding the neural control of head movements in different animal species. The superior colliculus (SC) has been identified as the primary brainstem center controlling the orienting. Its output signal is transmitted to neck motoneurons via two major separate pathways: one through the reticulospinal neurons (RSNs) in the pons and medulla and the other through neurons in Forel's field H (FFH) in the mesodiencephalic junction. The tecto-reticulo-spinal pathway controls orienting chiefly in the horizontal direction, while the tecto-FFH-spinal pathway controls orienting in the vertical direction. In each pathway, a subgroup of neurons functions as premotor neurons for both extraocular and neck motoneurons, while others are specified for each, which allows both coordinated and separate control of eye and head movements. Head movements almost always produce shifts in the center of gravity that might cause postural disturbances. The postural equilibrium may be maintained by transmitting the orienting command to the limb segments via descending axons of the reticulospinal and long propriospinal neurons. The SC and brainstem relay neurons receive descending inputs from higher order structures such as the cerebral cortex, cerebellum, and basal ganglia. These inputs may serve context-dependent control of orienting by modulating the activities of the primary brainstem pathways.

Animals↗

GABAB-receptor-mediated inhibition reduces the orientation selectivity of the sustained response of striate cortical neurons in cats.

Blocking GABAA-receptor-mediated inhibition reduces the selectivity of striate cortical neurons for the orientation of a light bar primarily by reducing the selectivity of their onset transient (initial 200 ms) response. Blocking GABAB-receptor-mediated inhibition with phaclofen, however, is not reported to reduce the orientation selectivity of these neurons when it is measured with a light bar. We hypothesized that blocking GABAB-receptor-mediated inhibition would instead affect the orientation selectivity of cortical neurons by reducing the selectivity of their sustained response to a prolonged stimulus. To test this hypothesis, we stimulated 21 striate cortical neurons with drifting sine-wave gratings and measured their orientation selectivity before, during, and after iontophoretic injection of 2-hydroxy-saclofen (2-OH-S), a selective GABAB-receptor antagonist. 2-OH-S reduced the orientation selectivity of six of eight simple cells by an average of 28.8 (+/- 13.2) % and reduced the orientation selectivity of eight of 13 complex cells by an average of 32.3 (+/- 27.4) %. As predicted, 2-OH-S reduced the orientation selectivity of the neurons' sustained response, but did not reduce the orientation selectivity of their onset transient response. 2-OH-S also increased the length of spike "bursts" (two or more spikes with interspike intervals < or = 8 ms) and eliminated the orientation selectivity of these bursts for six cells. These results are the first demonstration of a functional role for GABAB receptors in visual cortex and support the hypothesis that two GABA-mediated inhibitory mechanisms, one fast and the other slow, operate within the striate cortex to shape the response properties of individual neurons.

Action Potentials↗

Neuronal responses to orientation and motion contrast in cat striate cortex.

Responses of striate neurons to line textures were investigated in anesthetized and paralyzed adult cats. Light bars centered over the excitatory receptive field (RF) were presented with different texture surrounds composed of many similar bars. In two test series, responses of 169 neurons to textures with orientation contrast (surrounding bars orthogonal to the center bar) or motion contrast (surrounding bars moving opposite to the center bar) were compared to the responses to the corresponding uniform texture conditions (all lines parallel, coherent motion) and to the center bar alone. In the majority of neurons center bar responses were suppressed by the texture surrounds. Two main effects were found. Some neurons were generally suppressed by either texture surround. Other neurons were less suppressed by texture displaying orientation or motion (i.e. feature) contrast than by the respective uniform texture, so that their responses to orientation or motion contrast appeared to be relatively enhanced (preference for feature contrast). General suppression was obtained in 33% of neurons tested for orientation and in 19% of neurons tested for motion. Preference for orientation or motion contrast was obtained in 22% and 34% of the neurons, respectively, and was also seen in the mean response of the population. One hundred nineteen neurons were studied in both orientation and motion tests. General suppression was correlated across the orientation and motion dimension, but not preference for feature contrast. We also distinguished modulatory effects from end-zones and flanks using butterfly-configured texture patterns. Both regions contributed to the generally suppressive effects. Preference for orientation or motion contrast was not generated from either end-zones or flanks exclusively. Neurons with preference for feature contrast may form the physiological basis of the perceptual saliency of pop-out elements in line textures. If so, pop-out of motion and pop-out of orientation would be encoded in different pools of neurons at the level of striate cortex.

Animals↗

Mental representations of large and small spatial layouts are orientation dependent.

Previous research on spatial memory indicated that memories of small layouts were orientation dependent (orientation specific) but that memories of large layouts were orientation independent (orientation free). Two experiments investigated the relation between layout size and orientation dependency. Participants learned a small or a large 4-point path (Experiment 1) or a large display of objects (Experiment 2) and then made judgments of relative direction from imagined headings that were either the same as or different from the single studied orientation. Judgments were faster and more accurate when the imagined heading was the same as the studied orientation (i.e., aligned) than when the imagined heading differed from the studied orientation (i.e., misaligned). This alignment effect was present for both small and large layouts. These results indicate that location is encoded in an orientation-dependent manner regardless of layout size.

Female↗

Inhibitory processes in covert orienting in patients with Alzheimer's disease.

Previous studies of covert orienting in Alzheimer's disease (AD) have investigated exogenous and endogenous processes separately. We aimed to investigate how the 2 modes of orienting interact to control attention in healthy older participants and patients with AD. The covert orienting of visual attention task (COVAT) with abrupt onset cues was used in all experiments. In Experiments 1 and 2, predictive information was added to cues to initiate an endogenous orienting response. Results showed that healthy older participants were able to use endogenous processes to inhibit exogenous orienting. In contrast, patients with AD were unable to inhibit exogenous orienting to cues even when targets rarely appeared there. Experiment 3 investigated inhibition of return (IOR) in patients with AD. Both healthy older controls and patients with AD showed a normal IOR, suggesting that exogenous orienting processes are relatively unaffected by the normal aging process or in patients with AD. A model of covert orienting in which exogenous and endogenous orienting processes interact to control attentional behaviors is discussed.

Aged↗

Influence of experience on orientation maps in cat visual cortex.

Experience is known to affect the development of ocular dominance maps in visual cortex, but it has remained controversial whether orientation preference maps are similarly affected by limiting visual experience to a single orientation early in life. Here we used optical imaging based on intrinsic signals to show that the visual cortex of kittens reared in a striped environment responded to all orientations, but devoted up to twice as much surface area to the experienced orientation as the orthogonal one. This effect is due to an instructive role of visual experience whereby some neurons shift their orientation preferences toward the experienced orientation. Thus, although cortical orientation maps are remarkably rigid in the sense that orientations that have never been seen by the animal occupy a large portion of the cortical territory, visual experience can nevertheless alter neuronal responses to oriented contours.

Animals↗

Two types of orientation-sensitive responses of amacrine cells in the mammalian retina.

Neurons sensitive to the orientation of light stimuli exist throughout the mammalian visual system, suggesting that this spatial feature is a fundamental cue used by the brain to decipher visual information. The most peripheral neurons known to show orientation sensitivity are the retinal ganglion cells. Considerable morphological and pharmacological data suggest that the orientation sensitivity of ganglion cells is formed, at least partly, by the amacrine cells, which are laterally oriented interneurons presynaptic to the ganglion cells in the inner plexiform layer. So far there have been few studies of the responses of amacrine cells to oriented visual stimuli and their role in forming orientation-sensitive responses in the retina remains unclear. Here I report the novel finding of a population of amacrine cells in the rabbit retina which are orientation-sensitive. These amacrine cells can be divided into two subtypes, whose orientation sensitivity is manufactured by two distinct mechanisms. The orientation sensitivity of the first subtype of amacrine cell is formed from the interactions of excitatory, centre-receptive field synaptic inputs and inhibitory inputs of opposite polarity, whereas that for cells of the second subtype seems to be the product of a marked asymmetry in their dendritic arbors.

Animals↗

Evidence for a contribution of lateral inhibition to orientation tuning and direction selectivity in cat visual cortex: reversible inactivation of functionally characterized sites combined with neuroanatomical tracing techniques.

We have previously reported that cells in cat areas 17 and 18 can show increases in response to non-optimal orientations or directions, commensurate with a loss of inhibition, during inactivation of laterally remote, visuotopically corresponding sites by iontophoresis of gamma-aminobutyric acid (GABA). We now present anatomical evidence for inhibitory projections from inactivation sites to recording sites where 'disinhibitory' effects were elicited. We made microinjections of [3H]-nipecotic acid, which selectively exploits the GABA re-uptake mechanism, < 100 microm from recording sites where cells had shown either an increase in response to non-optimal orientations during inactivation of a cross-orientation site (n = 2) or an increase in response to the non-preferred direction during inactivation of an iso-orientation site with opposite direction preference (n = 5). Retrogradely labelled GABAergic neurons were detected autoradiographically and their distribution was reconstructed from series of horizontal sections. In every case, radiolabelled cells were found in the vicinity of the inactivation site (three to six within 150 microm). The injection and inactivation sites were located in layers II/III-IV and their horizontal separation ranged from 400 to 560 microm. In another experiment, iontophoresis of biocytin at an inactivation site in layer III labelled two large basket cells with terminals in close proximity to cross-orientation recording sites in layers II/III where disinhibitory effects on orientation tuning had been elicited. We argue that the inactivation of inhibitory projections from inactivation to recording sites made a major contribution to the observed effects by reducing the strength of inhibition during non-optimal stimulation in recurrently connected excitatory neurons presynaptic to a recorded cell. The results provide further evidence that cortical orientation tuning and direction selectivity are sharpened, respectively, by cross-orientation inhibition and iso-orientation inhibition between cells with opposite direction preferences.

Animals↗

Fast oscillations display sharper orientation tuning than slower components of the same recordings in striate cortex of the awake monkey.

We wanted to know whether fast oscillations ( approximately 30-80 Hz) in striate cortex of awake monkeys show sharper orientation selectivity than (i) slower components, including spike rate modulations, and (ii) broad-band signals of the same recordings. As fast oscillations are probably of cortical origin this may further clarify whether cortical network mechanisms are substantially involved in generating orientation selectivity. We recorded multi unit activity (MUA) and local field potentials (LFP, 1-140 Hz) by the same microelectrodes from upper layers of macaque striate cortex during visual stimulation with grating textures of different orientations. An orientation index (OI) was derived from the cortical responses in three frequency ranges (low, 0-11.7 Hz; medium, 11.7-31.3 Hz; and fast oscillations, 31.3-62.5 Hz) and for the broad-band LFP and MUA power. (i) Both LFP and MUA fast oscillations reveal a higher orientation index than signal components in the low and medium frequency ranges. (ii) For MUA the orientation index was significantly higher with fast oscillations than for the lower frequency ranges and the initial broad-band transient responses. (iii) LFPs show a significantly higher orientation index only for the fast oscillations during sustained activation compared with their broad-band power during the transient responses. Thus, our main result is the sharper orientation tuning of fast oscillations in spike activities of local populations compared with slower components of the same broad-band recordings. As fast oscillations occur synchronized in the awake monkey's striate cortex we assume that they have enhanced probability of activating successive stages of visual processing and hence contribute to the perception of orientation.

Animals↗

Edge computation in human vision: anisotropy in the combining of oriented filters.

Above threshold, two superimposed sinusoidal gratings of the same spatial frequency (eg 1 cycle deg-1) and equal contrasts, and with orientations balanced around vertical, usually look like a compound structure containing vertical and horizontal edges. However, at large plaid angles (ie large differences between component orientations) and low plaid contrasts there is a tendency for the stimulus to appear as two overlapping gratings (component structure) with obliquely oriented edges. These dependencies of perceived spatial structure in plaids are incompatible with an edge-coding scheme that uses only circular filters to compute zero-crossings, but instead support the idea that different oriented filters can (compound percept) or cannot (component percept) be combined before edges are represented. Here, further evidence is presented in support of this hypothesis. Two-component plaid stimuli had plaid angles of 45 degrees or 90 degrees, and a range of plaid orientations (ie a range of orientations around which the plaid components were balanced). Observers indicated whether each stimulus was perceived as a compound or component structure for a range of plaid contrasts. In addition to angle and contrast effects, perceived spatial structure was also found to depend on plaid orientation: compound structures were perceived more often when the plaid components were balanced around the cardinal axes of the retina. It is suggested that the principles governing the combination of oriented-filter outputs might be learnt during the development of the visual system by using a Hebb-type rule: coactivated filters are more likely to combine their outputs when activated on future occasions. Given the prominence of vertical and horizontal orientations in a carpentered environment, this simple rule promotes a network that combines filters balanced around cardinal axes more readily than oblique axes, in agreement with the results.

Anisotropy↗

Orientation priming of novel shapes in the context of viewpoint-dependent recognition.

Can visual similarity between shapes facilitate orientation priming? Five experiments are reported in which this possibility was explored by using novel two-dimensional shapes that formed homogeneous stimulus classes. After training on individual shapes in a canonical view, the recognition of these shapes was tested in several picture-plane orientations. In experiments 1 and 2 an identification task was used to replicate the classic finding obtained with the mirror-judgment task-that prior orientation cueing does not reduce the magnitude of orientation dependence in processing rotated shapes. The results of experiment 3, however, indicate that blocking trials by orientation is one condition in which orientation priming may be obtained. Experiment 4 builds on this result, and it is suggested that awareness of the blocking manipulation is not required to obtain orientation priming. In experiment 5 the mechanisms underlying this finding are explored, and evidence is offered that orientation priming is a consequence of representations that encode both shape and orientation. Such results may be considered as an extension to the 'image-based' approach to object recognition, demonstrating that generalization across exemplars may occur within recognition mechanisms that are viewpoint dependent.

Humans↗

Recognising the usual orientation of one's own face: the role of asymmetrically located details.

Our ability to recognise the usual horizontal orientation of our own face (mirror orientation) as compared with another very familiar face (normal orientation) was examined in experiment 1. Participants did not use the same kind of information in determining the orientation of their own face as in determining the orientation of the other familiar face. The proportion of participants who reported having based their judgment on the location of an asymmetric feature (eg a mole) was higher when determining the orientation of their own face than when determining that of the other familiar face. In experiment 2, participants were presented with pairs of manipulated images of their own face and of another familiar face showing conflicting asymmetric features and configural information. Each pair consisted of one picture showing asymmetric features of a given face in a mirror-reversed position, while the facial configuration was left unchanged; and one picture in which the location of the asymmetric features was left unchanged, while the facial configuration was mirror-reversed. As expected from the hypothesis that asymmetric local features are more frequently used for the judgment of one's own face, participants chose the picture showing mirror-reversed asymmetric features when determining the usual orientation of their own face significantly more often than they chose the picture showing normally oriented asymmetric features when determining the orientation of the other face. These results are explained in terms of competing forward and mirror-reversed representations of one's own face.

Adult↗

The effect of text orientation, visual meridian, and inter-character spacing on word identification in the retinal periphery.

Previous research has demonstrated that the masking effects of flankers about a target in the peripheral retina are not isotropic. Rather, regions of lateral interaction are ellipsoid in shape with the major axis oriented radially along a meridian through the fovea. This finding leads to the counterintuitive prediction that horizontal text positioned to the right of fixation might be read more slowly than similarly positioned text oriented diagonally or vertically. Similarly, vertically oriented text above fixation might be read more slowly than horizontally or diagonally oriented text above fixation. We investigated the effect of text orientation and inter-character spacing on word identification in the retinal periphery. Text was presented by rapid serial visual presentation. Words were centered 3 degrees from fixation along four visual field meridians (VM) (right horizontal, upper-right diagonal, vertical, and upper-left diagonal). Regardless of VM identification, performance was best for horizontal text, declining slightly for orientations between +60 degrees and -60 degrees and declining more quickly for acute orientations. A weak effect of VM was observed for text with normal inter-character spacing. Performance was best for text centered along the horizontal meridian and declined slightly along the other VM. Finally, identification rates increased by approximately 33 words min(-1) with the addition of one character space between adjacent letters. The word-recognition processes are very tolerant of text orientation, exhibiting a modest decline for orientations within +/- 60 of horizontal regardless of VM.

Adult↗