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PubMed1966

Orientational selectivity of the human visual system.

1. It is known that an object is less detectable when it is viewed against a background containing structures similar to the object. The effect of changing the orientation between the object and background is investigated.2. Gratings of variable contrast were generated on two oscilloscopes; these were superimposed optically. The angle of orientation between them could be changed. The threshold of one grating, the test grating, was determined in the presence of the other, the masking grating.3. When the gratings were presented with the same orientation (and locked in phase) the increment threshold of the test grating was found to be proportional to the suprathreshold contrast of the masking grating.4. As the angle between the test and masking gratings was increased the masking effect fell exponentially.5. At 12 degrees on either side of a vertical test grating the masking effect was reduced by a factor of two with respect to its maximum value. This angle was independent of the contrast level of masking, the focus, and also the phase coherence of the masking grating.6. If the test grating was presented obliquely the effect of masking was slightly less.7. The narrow orientationally tuned channels found psychophysically by this masking technique are compared with the orientationally sensitive cells discovered electrophysiologically in the visual cortex of the cat.

Cerebral Cortex↗
PubMed1977

Spectral and orientation specificity of single cells in foveal striate cortex of the vervet monkey, Cercopithecus aethiops.

1. The spectral sensitivity, orientation specificity and inhibitory surround of seventy-three cells were studied in the vervet monkey. The eye was in the dark or illuminated with steady white or spectral light. The cells were in the striate cortex corresponding to the foveal representation. Nearly all the cells gave on- or on/off-responses.2. More than a third of the cells (41%) responded over a narrow spectral band, in the blue, green, yellow or red section. Three quarters of them were orientation specific with flanking inhibitory surround and half of these were of the ;stopped-end' variety as well. The effect of the wave-length of the background indicated that only a half were activated by such excitatory-inhibitory colour pairs as have been described in the geniculate nucleus.3. A third of the cells (36%) responded to most colours but with the greatest response to green, yellow or red. Less than half were orientation specific. Unlike the narrowband cells, the response decreased with the intensity of the light. Two thirds were activated by the excitatory-inhibitory colour pairs that have been described in the geniculate nucleus.4. In both groups of cells the wave-length of the spectral peak could be different when the stimuli were weak compared with when they were strong.5. One quarter of the cells (23%) gave on/off-responses to all spectral flashes; half were not orientation specific. The difference in orientation specificity between narrow- and broadband cells is significant (P < 0.01).

Action Potentials↗
PubMed2002

Dynamics and constancy in cortical spatiotemporal patterns of orientation processing.

How does the high selectivity to stimulus orientation emerge in the visual cortex? Thalamic feedforward-dominated models of orientation selectivity predict constant selectivity during the visual response, whereas intracortical recurrent models predict dynamic improvement in selectivity. We imaged the cat visual cortex with voltage-sensitive dyes to measure orientation-tuning dynamics of a large neuronal population. Tuning-curve width did not narrow after response onset, whereas the difference between preferred and orthogonal responses (modulation depth) first increased, then declined. We identified a suppression of the evoked responses, referred to as the evoked deceleration-acceleration (DA) notch, which was larger for the orthogonal response. Furthermore, peak selectivity of the tuning curves was contemporaneous with the evoked DA notch. These findings suggest that in the cat brain, sustained visual cortical processing does not narrow orientation tuning; rather, intracortical interactions may amplify modulation depth and suppress the orthogonal response relatively more than the preferred. Thus, feedforward models and recurrent models of orientation selectivity must be combined.

Animals↗
PubMed2000

The contribution of noise to contrast invariance of orientation tuning in cat visual cortex.

Feedforward models of visual cortex appear to be inconsistent with a well-known property of cortical cells: contrast invariance of orientation tuning. The models' fixed threshold broadens orientation tuning as contrast increases, whereas in real cells tuning width is invariant with contrast. We have compared the orientation tuning of spike and membrane potential responses in single cells. Both are contrast invariant, yet a threshold-linear relation applied to the membrane potential accurately predicts the orientation tuning of spike responses. The key to this apparent paradox lies in the noisiness of the membrane potential. Responses that are subthreshold on average are still capable of generating spikes on individual trials. Unlike the iceberg effect, contrast invariance remains intact even as threshold narrows orientation selectivity. Noise may, by extension, smooth the average relation between membrane potential and spike rate throughout the brain.

Action Potentials↗
PubMed2001

A recurrent neural network for partitioning of hand drawn characters into strokes of different orientations.

A neural network is described which is intended to extract orientation features that should be used for recognition of hand drawn characters. The network partitions the input hand drawn characters into separate line segments (strokes) according to their orientations. The network consists of several neural layers; each layer serves for extracting strokes of a certain orientation. Every neural layer has one-to-one correspondence with an input screen. The network uses an iterative update procedure which includes interactions of neurons inside each layer through oriented excitatory connections and inhibitory interrelations between the corresponding neurons of different layers. Computer simulation of the network was performed. Experiments showed that the network efficiently classifies all pixels of any hand drawn characters according to the orientations of the strokes constituting these characters and performs, as a result of that, a reasonable segmentation of characters.

Algorithms↗
PubMed2003

Simultaneous control of hand displacements and rotations in orientation-matching experiments.

In reach-to-grasp movements, the interaction between the hand changes in position and those in orientation is poorly understood. A theoretical approach previously proposed (Torres EB and Zipser D. J Neurophysiol 88: 1-13, 2002) assumes that motion strategies are resolved in space independently from the temporal dynamics of the motion and predicts the coarticulation of the hand transport and rotation along the path. The model implies that this simultaneous control is independent of variations in speed and initial posture and required matching orientation. This paper presents experimental data from human subjects that confirm the model's predictions in the context of realistic, unconstrained, orientation-matching motions. Speed independence is quantified in the similarity of the postural and endpoint position-orientation paths obtained under three different speeds. Significant differences in hand and joint kinematics are shown in response to changes in initial posture and target orientation. The robustness of coarticulation under all three experimental conditions supports the idea of an intermediate stage that resolves the geometry of the motion independent of its temporal dynamics.

Arm↗
PubMed2005

Orientation-selective adaptation to first- and second-order patterns in human visual cortex.

Second-order textures-patterns that cannot be detected by mechanisms sensitive only to luminance changes-are ubiquitous in visual scenes, but the neuronal mechanisms mediating perception of such stimuli are not well understood. We used an adaptation protocol to measure neural activity in the human brain selective for the orientation of second-order textures. Functional MRI (fMRI) responses were measured in three subjects to presentations of first- and second-order probe gratings after adapting to a high-contrast first- or second-order grating that was either parallel or orthogonal to the probe gratings. First-order (LM) stimuli were generated by modulating the stimulus luminance. Second-order stimuli were generated by modulating the contrast (CM) or orientation (OM) of a first-order carrier. We used four combinations of adapter and probe stimuli: LM:LM, CM:CM, OM:OM, and LM:OM. The fourth condition tested for cross-modal adaptation with first-order adapter and second-order probe stimuli. Attention was diverted from the stimulus by a demanding task at fixation. Both first- and second-order stimuli elicited orientation-selective adaptation in multiple cortical visual areas, including V1, V2, V3, V3A/B, a newly identified visual area anterior to dorsal V3 that we have termed LO1, hV4, and VO1. For first-order stimuli (condition LM:LM), the adaptation was no larger in extrastriate areas than in V1, implying that the orientation-selective first-order (luminance) adaptation originated in V1. For second-order stimuli (conditions CM:CM and OM:OM), the magnitude of adaptation, relative to the absolute response magnitude, was significantly larger in VO1 (and for condition CM:CM, also in V3A/B and LO1) than in V1, suggesting that second-order stimulus orientation was extracted by additional processing after V1. There was little difference in the amplitude of adaptation between the second-order conditions. No consistent effect of adaptation was found in the cross-modal condition LM:OM, in agreement with psychophysical evidence for weak interactions between first- and second-order stimuli and computational models of separate mechanisms for first- and second-order visual processing.

Adaptation, Physiological↗
PubMed2003

From another angle: Differences in cortical coding between fine and coarse discrimination of orientation.

We measured the information available for orientation discrimination from metric distances for 24 cells in area 17 of cats that were paralyzed and anesthetized with Propofol and N(2)O. The metric distance information confirms fundamental coding differences for discrimination between fine (<10 degrees ) and coarse (>10 degrees ) orientation differences. The information for discriminating larger orientation differences is contained mainly in the firing rate, with minor enhancements from the coarse (30-70 ms) temporal structure in the firing rate. Both precise spike timing (9.2 ms) and intervals (6.8 ms) sustained over the stimulus presentation provide information for fine discrimination of orientation, where almost no reliable information is provided by the spike count. We compare and confirm the results (using the same data set) to vector distances based on classification theory. The results support a dynamic spiking mechanism where coordinated activity could provide fast and reliable information about detailed angle and/or direction information in the region of the preferred orientation.

Animals↗
PubMed1987

Linear mechanism of orientation tuning in the retina and lateral geniculate nucleus of the cat.

1. The orientation tuning of lateral geniculate nucleus (LGN) neurons and retinal ganglion cells (recorded as S potentials in the LGN) was investigated with drifting grating stimuli. 2. Results were compared with a quantitative model, in which receptive fields were constructed from linear, elliptical Gaussian center and surround subunits, and responses could be predicted to gratings of any spatial frequency at any orientation. 3. The orientation tuning of X and Y retinal ganglion cells and LGN neurons was shown to result from the linear mechanism of receptive-field elongation, as data from these cells could be well fit with this model. 4. The responses of LGN neurons and their input retinal ganglion cells were compared. The orientation tuning of LGN neurons was found to be a reflection of the tuning of their retinal inputs, showing that neither intrageniculate neural interactions nor the corticogeniculate projection play any role in LGN orientation selectivity.

Action Potentials↗
PubMed1990

Selectivity for orientation and direction of motion of single neurons in cat striate and extrastriate visual cortex.

1. We consider the consequences of the orientation selectivity shown by most cortical neurons for the nature of the signals they can convey about the direction of stimulus movement. On theoretical grounds we distinguish component direction selectivity, in which cells are selective for the direction of movement of oriented components of a complex stimulus, from pattern direction selectivity, or selectivity for the overall direction of movement of a pattern irrespective of the directions of its components. We employed a novel test using grating and plaid targets to distinguish these forms of direction selectivity. 2. We studied the responses of 280 cells from the striate cortex and 107 cells from the lateral suprasylvian cortex (LS) to single sinusoidal gratings to determine their orientation preference and directional selectivity. We tested 73 of these with sinusoidal plaids, composed of two sinusoidal gratings at different orientations, to study the organization of the directional mechanisms within the receptive field. 3. When tested with single gratings, the directional tuning of 277 oriented cells in area 17 had a mean half width of 20.6 degrees, a mode near 13 degrees, and a range of 3.8-58 degrees. Simple cells were slightly more narrowly tuned than complex cells. The selectivity of LS neurons for the direction of moving gratings is not markedly different from that of neurons in area 17. The mean direction half width was 20.7 degrees. 4. We evaluated the directional selectivity of these neurons by comparing responses to stimuli moved in the optimal direction with those elicited by a stimulus moving in the opposite direction. In area 17 about two-thirds of the neurons responded less than half as well to the non-preferred direction as to the preferred direction; two-fifths of the units responded less than one-fifth as well. Complex cells showed a somewhat greater tendency to directional bias than simple cells. LS neurons tended to have stronger directional asymmetries in their response to moving gratings: 83% of LS neurons showed a significant directional asymmetry. 5. Neurons in both areas responded independently to each component of the plaid. Thus cells giving single-lobed directional-tuning curves to gratings showed bilobed plaid tuning curves, with each lobe corresponding to movement in an effective direction by one of the two component gratings within the plaid. The two best directions for the plaids were those at which one or other single grating would have produced an optimal response when presented alone.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Population encoding of spatial frequency, orientation, and color in macaque V1.

1. We recorded local field potentials in the parafoveal representation in the primary visual cortex of anesthetized and paralyzed macaque monkeys with a multicontact electrode that provided for sampling of neural activity at 16 sites along a vertical penetration. Differential recordings at adjacent contacts were transformed into an estimate of current source density (CSD), to provide a measure of local neural activity. 2. We used m-sequence stimuli to map the region of visual space that provided input to the recording site. The local field potential recorded in macaque V1 has a population receptive field (PRF) size of approximately 2 deg2. 3. We assessed spatial tuning by the responses to two-dimensional Gaussian noise, spatially filtered to retain power only within one octave. Responses to achromatic band-limited noise stimuli revealed a prominent band-pass spatial tuning in the upper layers, but a more low-pass spatial tuning in lower layers. 4. We assessed orientation tuning by the responses to band-limited noise whose spectrum was further restricted to lie within 45 degrees wedges. The local field potential showed evidence of orientation tuning at most sites. Orientation tuning in upper and lower layers was manifest by systematic variations not only in response size but also in response dynamics. 5. We assessed chromatic tuning by the responses to isotropic band-limited noise modulated in a variety of directions in tristimulus space. Some lower-layer locations showed a nulling of response under near-isoluminant conditions. However, response dynamics in upper and lower layers depended not only on luminance contrast, but also on chromatic inputs. 6. Responses to near-isoluminant stimuli and to low-contrast luminance modulation were shifted to lower spatial frequencies. 7. We determined the extent to which various temporal frequencies in the response conveyed information concerning spatial frequency, orientation, and color under the steady-state conditions used in these studies. In each case, information is distributed in the response dynamics across a broad temporal frequency range, beginning at 4 Hz (the lowest frequency used). For spatial frequency the information rate remains significant up to at least 25 Hz. For orientation tuning and chromatic tuning, the information rate is lower overall and remains significant up to 13 Hz. In contrast, for texture discrimination, information is shifted to lower temporal frequencies.

Animals↗
PubMed1995

Activity of neurons in the medial pontomedullary reticular formation during orienting movements in alert head-free cats.

1. Single unit activities of 236 neurons were recorded in the medial pontomedullary reticular formation during visually triggered orienting gaze shifts in 10 alert cats under head-free conditions using movable tungsten-needle electrodes attached to the skull. The activities were analyzed mainly in relation to the head movement that was triggered by presentation of a light-emitting diode (LED) in one of eight directions separated radially by 45 deg after fixation of the center LED. Of these, 120 neurons were recorded in the pontine reticular formation, chiefly in the nucleus reticularis pontis caudalis, and the remaining 116 were in the medullary reticular formation, chiefly in the nucleus reticularis gigantocellularis. Activities of 65 pontine and 65 medullary neurons were modulated in relation to the dynamic phase of orienting movements ("orienting-related neurons"). Activities of the remaining neurons were modulated either irregularly or not at all during orienting movement ("irregular or no-response neurons"). Input from the contralateral superior colliculus and cerebral cortex and projections to the spinal cord were also investigated. 2. Among the orienting-related neurons, 62 pontine and 55 medullary neurons showed increases in activity preceding the onset of eye and head movement by 0-155 ms ("pretype"). Three pontine and 10 medullary neurons showed increases in activity only after the onset of movement ("posttype"). Of the pretype neurons, 61 pontine and 51 medullary neurons showed directional preference of activity ("directional" neurons). One pontine and four medullary neurons were classified as "omnidirectional" because these neurons increased activity preceding movements in all directions tested, and no directional preference was apparent. 3. In the pretype-directional cells, the average firing frequency during bursts was correlated with amplitude and angular velocity of head movements. Activities of the directional neurons during movements in the eight different directions could be well fitted with cosine functions in the majority of cases. The preferred directions of most pontine neurons and of about half the medullary neurons, as determined by first-degree sinusoidal regression analysis, were distributed around the ipsiversive horizontal axis. However, there were also a considerable number of neurons whose preferred directions were upward, downward, contraversive, or oblique in the medulla. 4. Among the directional cells preferring ipsiversive horizontal movements, 11 pontine neurons showed activity, the onset of which was locked to visual stimuli with latencies of 40-70 ms, in addition to phasic discharges locked to the onset of movement. This "stimulus-locked activity" was sometimes modulated depending on the attentional state of the animal.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1997

Spatial and temporal integration of signals in foveal line orientation.

The discrimination of the orientation of a line improves with line length, reaching an optimum when a foveal line is approximately 0.5 degrees long. We studied the effect of eliminating sections of the line, of displacing them out of alignment, and of delaying them. Orientation discrimination thresholds are only a little elevated when a 25-arcmin line is replaced by three equally spaced collinear 5-arcmin segments. Two collinear 5-arcmin segments show better thresholds than a single one when they are separated by as much as 20 arcmin. But thresholds are impaired by bringing line segments out of collinearity by as little as 1 arcmin. Asynchrony of up to 50 ms can be tolerated, but when the middle segment of a three-line pattern is delayed by approximately 100 ms there is active inhibition, thresholds being now higher than when the middle segment is absent. It is concluded that for signals to address the orientation discrimination mechanism optimally, they have to be contained inside a narrow spatial corridor and be presented within a time window of approximatley 50 ms, but that some spatial summation can take place over a length of > or = 0.5 degrees in the fovea. Because short lines made of black and white collinear segments do not have good orientation thresholds, whereas longer and interrupted lines do, it is concluded that what is involved is potentiating interaction between collinearly arranged neurons with identical orientation selectivity rather than summation of signals within the receptive fields of single neurons.

Adult↗
PubMed2000

Discrimination of line orientation in humans and monkeys.

Orientation discrimination, the capacity to recognize an orientation difference between two lines presented at different times, probably involves cortical processes such as stimuli encoding, holding them in memory, comparing them, and then deciding. To correlate discrimination with neural activity in combined psychophysical and electrophysiological experiments, precise knowledge of the strategies followed in the completion of the behavioral task is necessary. To address this issue, we measured human and nonhuman primates' capacities to discriminate the orientation of lines in a fixed and in a continuous variable task. Subjects have to indicate whether a line (test) was oriented to one side or to the other of a previously presented line (reference). When the orientation of the reference line did not change across trials (fixed discrimination task), subjects can complete the task either by categorizing the test line, thus ignoring the reference, or by discriminating between them. This ambiguity was avoided when the reference stimulus was changed randomly from trial to trial (continuous discrimination task), forcing humans and monkeys to discriminate by paying continuous attention to the reference and test stimuli. Both humans and monkeys discriminated accurately with stimulus duration as short as 150 ms. Effective interstimulus intervals were of 2.5 s for monkeys but much longer (>6 s) in humans. These results indicated that the fixed and continuous discrimination tasks are different, and accordingly humans and monkeys do use different behavioral strategies to complete each task. Because both tasks might involve different neural processes, these findings have important implications for studying the neural mechanisms underlying visual discrimination.

Adolescent↗
PubMed2002

Color processing in macaque striate cortex: relationships to ocular dominance, cytochrome oxidase, and orientation.

We located clusters of color-selective neurons in macaque striate cortex, as mapped with optical imaging and confirmed with electrophysiological recordings. By comparing responses to an equiluminant red/green stimulus versus a high-contrast luminance stimulus, we were able to reveal a patchy distribution of color selectivity. Other color imaging protocols, when compared with electrophysiological data, did not reliably indicate the location of functional structures. The imaged color patches were compared with other known functional subdivisions of striate cortex. There was a high degree of overlap of the color patches with the cytochrome-oxidase (CO) blobs. The patches were often larger than a single blob in size, however, and in some instances spanned two neighboring blobs. More than one-half (56%) of the color-selective patches seen in optical imaging were not confined to one ocular dominance (OD) column. Almost one-quarter of color patches (23%) extended across OD columns to encompass two blobs of different eye preference. We also compared optical images of orientation selectivity to maps of color selectivity. Results indicate that the layout of orientation and color selectivity are not directly related. Specifically, despite having similar scales and distributions, the maps of orientation and color selectivity were not in consistent alignment or registration. Further, we find that the maps of color selectivity and of orientation are each only loosely related to maps of OD. This description stands in contrast to a common depiction of color-selective regions as identical to CO blobs, appearing as pegs in the centers of OD columns in the classical "ice cube" model. These results concerning the pattern of color selectivity in V1 support the view (put forth in previous imaging studies of the organization of orientation and ocular dominance) that there is not a fundamental registration of functional hypercolumns in V1.

Animals↗
PubMed1998

The effects of telencephalic lesions on visually mediated prey orienting behavior in the leopard frog (Rana pipiens). II. The effects of limited lesions to the telencephalon.

Unilateral removal of the telencephalon in the leopard frog, Rana pipiens, produces a contralateral deficit in visual prey orienting behavior [Patton and Grobstein, 1997]. In mammals, such deficits are most commonly associated with damage to the isocortex, a pallial derived structure. In contrast, we here report that in leopard frogs, lesions that remove substantial areas of one telencephalic lobe, including virtually the entire pallium, have no discernible effect on visual orienting behavior. Restricted lesions to the ventrocaudal telencephalon, however, produce an effect that closely resembles that produced by the complete removal of one telencephalic lobe. The 'critical area' that is both included in all lesions that are effective in producing a severe deficit and excluded from all ineffective lesions includes a portion of the caudal striatum. The striatum is known to play a significant role in anuran vision. It thus seems likely that the deficit produced by unilateral removal of the telencephalon in the leopard frog is due specifically to the removal of the caudal striatum. Unilateral lesions to the striatum have previously been shown to produce a contralateral deficit in visual orienting behavior in cats, and a role for the striatonigral pathway in the production of the visual orienting deficit that follows visual cortex lesions has been proposed. The current findings call attention to the possible general importance of the striatum in the control of vertebrate visual orienting behaviors.

Animals↗
PubMed2003

Interocular orientation disparity and the stereoscopic perception of slanted surfaces.

The orientation threshold for two-dimensional filtered noise stimuli was estimated using forced-choice procedures with both dioptic and dichoptic viewing. In the dioptic case the two patterns were co-rotated. In the dichoptic case the stimuli were counter-rotated to produce an orientation disparity, which yields a percept of slant about the horizontal axis orthogonal to the cyclopean line of sight. Dioptic thresholds increased with the orientation bandwidth of the stimuli. In contrast, dichoptic thresholds were essentially constant across a wide range of conditions. In all cases, dichoptic orientation acuity was much finer than conventional estimates. In a second experiment, the dichoptic threshold was estimated for patterns superimposed on a depth pedestal. Acuity was affected significantly by the presence of the pedestal, and was an inverse function of pedestal amplitude. The results suggest that stereoscopic slant caused by dichoptic counter-rotation arises because of neural processing of the overall pattern of disparities of position produced by counter-rotation, rather than specialised encoding of orientation disparity.

Depth Perception↗
PubMed2003

Interactions between color and luminance in the perception of orientation.

At the early stages of visual processing in humans and other primates, chromatic signals are carried to primary visual cortex (V1) via two chromatic channels and a third achromatic (luminance) channel. The sensitivities of the channels define the three cardinal axes of color space. A long-standing though controversial hypothesis is that the cortical pathways for color and form perception maintain this early segregation with the luminance channel dominating form perception and the chromatic channels driving color perception. Here we show that a simple interaction between orientation channels (the tilt illusion) is influenced by both chromatic and luminance mechanisms. We measured the effect of oriented surround gratings upon the perceived orientation of a test grating as a function of the axes of color space along which the gratings were modulated. We found that the effect of a surround stimulus on the perceived orientation of the test is largest when both are modulated along the same axis of color space, regardless of whether that is a cardinal axis. These results show that color and orientation are intimately coupled in visual processing. Further, they suggest that the cardinal chromatic axes have no special status at the level(s) of visual cortex at which the tilt illusion is mediated.

Color Perception↗