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PubMed1997

How Chlamydomonas keeps track of the light once it has reached the right phototactic orientation.

By using a real-time assay that allows measurement of the phototactic orientation of the unicellular alga Chlamydomonas with millisecond time resolution, it can be shown that single photons not only induce transient direction changes but that fluence rates as low as 1 photon cell(-1) s(-1) can already lead to a persistent orientation. Orientation is a binary variable, i.e., in a partially oriented population some organisms are fully oriented while the rest are still at random. Action spectra reveal that the response to a pulsed stimulus follows the Dartnall-nomogram for a rhodopsin while the response to a persistent stimulus falls off more rapidly toward the red end of the spectrum. Thus light of 540 nm, for which chlamy-rhodopsin is equally sensitive as for 440-nm light, induces no measurable persistent orientation while 440-nm light does. A model is presented which explains not only this behavior, but also how Chlamydomonas can track the light direction and switches between a positive and negative phototaxis. According to the model the ability to detect the direction of light, to make the right turn and to stay oriented, is a direct consequence of the helical path of the organism, the orientation of its eyespot relative to the helix-axis, and the special shielding properties of eyespot and cell body. The model places particular emphasis on the fact that prolonged swimming into the correct direction not only requires making a correct turn initially, but also avoiding further turns once the right direction has been reached.

Animals↗
PubMed1997

Evidence of visual processing negativity with attention to orientation and color in central space.

The aim of this study was to determine whether the visual frontal processing negativity reported in our earlier paper (Karayanidis, F. and Michie, P.T. Electroenceph. clin. Neurophysiol., 1996, 99: 38-56) is related to selection of spatial location, or occurs regardless of the stimulus features used to define the target. Subjects were instructed to respond to infrequent target stimuli of a particular combination of orientation, color and size. All stimuli were presented at central fixation. Posteriorly, orientation selection enhanced P125 amplitude over the right hemisphere but neither orientation nor color selection had an effect on N190. Posterior selection negativities emerged for orientation, color and their conjunction. At anterior sites, widespread effects of orientation and color processing were evident. The effect of location selection on the anterior N1 seen in our previous study was not evident with orientation selection. Instead, selection of orientation, color and their conjunction resulted in P145-250 frontally. Two later anterior negativities emerged. The early negativity (vPNe) was affected independently by orientation and color selection while the late negativity (vPNl) was affected only by selection of feature conjunction. Thus, the present results show that, like its auditory counterpart, the visual processing negativity occurs with a variety of stimulus classification features and is not exclusively related to spatial selection.

Adolescent↗
PubMed1997

Relating the mechanisms of orienting and alerting.

Cues provide two types of information: information about where the target will occur and when it will occur. We hypothesized two underlying processes related to cues, orienting (to location) and alerting. Using a covert orienting task under different conditions of alertness, we found evidence of independence between orienting and alerting (Experiments 3-4). The alerting mechanism is spatially broad and seems common for auditory and visual input (Experiments 1-2). In Experiment 1, visual cues at four locations occur simultaneously to prevent orienting; response facilitation was the same for targets occurring near or far from a cue. In Experiment 2, adding a visual alerting signal to an auditory signal provided no additional benefit. In Experiment 3, an auditory signal was used to modulate the alertness level during a covert orienting task. Orienting, measured by the validity effect, was independent of the level of alertness in this simple reaction task. Experiment 4 extended those results to a choice task. These studies indicate separate mechanisms of alerting and orienting. The global mode of alertness is consistent with the broad axonal distribution of the noradrenergic system. In contrast, human and animal data suggest that the orienting mechanism may be modulated by the basal forebrain cholinergic system.

Acoustic Stimulation↗
PubMed1999

Multimodal spatial orientation deficits in left-sided visual neglect.

Patients with right-sided temporo-parietal lesions often show contralesional neglect. However, neglect patients may also show spatial-perceptual deficits beyond the bisection and space exploration deficits frequently assessed in the horizontal plane, that is, deficits in the judgment of the subjective visual vertical or horizontal. In a recent study (Kerkhoff, G. & Zoelch, C.. Disorders of visuo-spatial orientation in the frontal plane in patients with visual neglect following right or left parietal lesions. Exp. Brain Res., 1998;122:108-120) we found significant perturbations in the perception of these three visual spatial axes in patients with contralesional neglect from right or left parietal lesions. To examine if this finding extends also to another modality we investigated how neglect patients perform tasks of visual- and tactile-spatial judgments of axis-orientation in the frontal plane. Visual-spatial and tactile-spatial judgments of the subjective vertical, horizontal and a right oblique orientation were obtained from patients with and without neglect as well as from normal subjects. Patients with left neglect showed a significant, contraversive tilt of all three visual-spatial axes (+5.6 degrees to +9.5 degrees, counterclockwise), and of the three tactile-spatial axes as well (+5.2 degrees to +10.5 degrees, counterclockwise). In contrast, right and left hemisphere lesioned control patients without neglect and normal control subjects showed unimpaired visual and tactile-spatial judgments (constant errors: < 1.0 degree). Difference thresholds in the visual-spatial tasks and unsigned errors in the tactile-spatial tasks were selectively elevated in the neglect group in contrast to all other subject groups. Spatial orientation deficits were significantly associated with the severity of clinical neglect (r = 0.55-0.88), and with the patients' ambulation performance (r = 0.45-0.70). Furthermore, crossmodal axis orientation tests in two neglect patients showed a similar counterclockwise tilt of +5 degrees to +15 degrees, suggesting a similar spatial deficit in both modalities. Orientation judgments were significantly aggravated by a 25 degree-tilt of the head to the left, as tested in one neglect patient, while a comparable rightward head-tilt improved spatial judgments in both modalities. This suggests that spatial orientation judgments are significantly modulated by gravitational input in neglect patients. Together these results are interpreted as evidence for multisensory spatial orientation deficits in neglect patients which are modulated by head-position and are related to their accompanying postural impairment.

Adult↗
PubMed2001

The effect of orientation learning on contrast sensitivity.

Regan and Beverley [Regan, D., & Beverley, K. I. (1985). Postadaptation orientation discrimination. Journal of the Optical Society of America A, 2(2), 147-155] previously demonstrated that adapting to an oriented visual stimulus improves sensitivity to subtle orientation differences while impairing contrast sensitivity. Here, we investigated whether practice-based improvements in orientation sensitivity would, like adaptation, impair contrast sensitivity. To the contrary, we found that contrast sensitivity actually improved significantly after observers demonstrated practice-based increases in orientation sensitivity. Therefore, while orientation sensitivity can be enhanced either by orientation-discrimination training or by adapting to visual stimuli, these two procedures have opposite effects on contrast sensitivity. This difference suggests that adaptation and perceptual learning on orientation discrimination cannot be explained sufficiently by a shared underlying cause, such as a reduction in neural activity.

Adaptation, Physiological↗
PubMed2001

Interaction between first- and second-order orientation channels revealed by the tilt illusion: psychophysics and computational modelling.

This paper examines the interaction between first- and second-order contours in the orientation domain. Using the simultaneous tilt illusion (TI), we show that the apparent rotation of a vertical test grating away from that of a surrounding inducing grating (repulsion effect) occurs when both the inducing and test grating are either first- or second-order. Furthermore, a significant repulsion effect is obtained when a first-order inducing grating surrounds a second-order test. If lateral inhibitory interactions between populations of orientation selective neurons provides a plausible explanation for orientation repulsion effects [Blakemore, C. B. Carpenter, R. H. S. & Georgeson, M. A. (1970) Nature, 228, 37-39], it is likely that the cue-invariant mechanisms that encodes the orientation of first- and second-order contours also exhibit inhibitory interactions. A two-channel computational model of orientation encoding is presented where one channel encodes only first-order stimuli while the second channel encodes both first- and second-order contours. In addition to predicting the orientation repulsion effects we observed, the model also provides a functional account of orientation attraction effects in terms of the responses of populations of orientation-tuned neurons.

Chi-Square Distribution↗
PubMed2002

Spatiotemporal interactions in detection of texture orientation modulations.

Previous studies have revealed spatial and temporal characteristics of texture orientation modulation detection. This study examined spatiotemporal interactions. We measured threshold amplitudes for detecting orientation modulations in various waveforms. The orientation modulations were presented in a dynamic texture display in which the spatial arrangement and mean orientation of elements were randomly updated at a given frame duration (17-900 ms). The results of three experiments all indicated significant spatiotemporal interactions. As the frame duration was decreased, the detection sensitivity declined more steeply for the sinusoidal orientation modulations than for the square and missing-fundamental waveforms (Expt 1), declined more steeply for low spatial-frequency sinusoidal modulations than for high frequency ones (Expt 2), and declined more steeply for sparse textures than for dense textures (Expt 3). These results indicate that the visual system loses its sensitivity more profoundly for long-range orientation modulations than for short-range modulations as the rate of orientation change increases, suggesting that the mechanism for detecting orientation modulation reduces its effective spatial range for rapid input changes.

Contrast Sensitivity↗
PubMed2003

Orientation opponency in human vision revealed by energy-frequency analysis.

Studies of second-order visual processing have primarily been concerned with understanding the mechanisms for detecting spatiotemporal variations in such attributes as contrast, orientation, spatial frequency, etc. Here, we have examined the orientation characteristics of second-order processes using bandpass noise whose Fourier energy is sinusoidally modulated across orientation, rather than across space or time. Sensitivity for detecting orientation-energy modulations was measured as a function of modulation frequency. The sensitivity function was bandpass, with a pronounced peak at an orientation frequency of 4 cycles/pi. An inverse Fourier transform of the sensitivity function revealed a filter profile displaying a centre-surround antagonism across orientation, with an excitatory centre within 6-9 deg and inhibitory lobes at 15-20 deg from the filter's centre. The degree of centre-surround antagonism increased with stimulus size far beyond the spatial range of the first-order filters (more than 64 times the dominant spatial wavelength of the noise carrier). These results suggest that second-order processing involves 'orientation-opponent' channels that extract differences in first-order outputs across orientation over a wide area of the visual field.

Contrast Sensitivity↗
PubMed2000

Adaptation-induced plasticity of orientation tuning in adult visual cortex.

A key emergent property of the primary visual cortex (V1) is the orientation selectivity of its neurons. The extent to which adult visual cortical neurons can exhibit changes in orientation selectivity is unknown. Here we use single-unit recording and intrinsic signal imaging in V1 of adult cats to demonstrate systematic repulsive shifts in orientation preference following short-term exposure (adaptation) to one stimulus orientation. In contrast to the common view of adaptation as a passive process by which responses around the adapting orientation are reduced, we show that changes in orientation tuning also occur due to response increases at orientations away from the adapting stimulus. Adaptation-induced orientation plasticity is thus an active time-dependent process that involves network interactions and includes both response depression and enhancement.

Adaptation, Physiological↗
PubMed1990

Bar orientation discrimination in the cat.

We have measured orientation-discrimination thresholds of 4 deg in the cat, confirming an earlier study of Vandenbussche and Orban (1983). Unlike Vandenbussche and Orban (1983), we found that orientation-discrimination performance is not better at principal, as compared to oblique, reference orientations (no oblique effect). Despite the absence of the oblique effect, and despite the discrimination thresholds which were elevated by a factor of 4 compared to humans, orientation-discrimination performance of cats and humans is qualitatively similar in a number of aspects. First, orientation-discrimination performance as a function of length and contrast is qualitatively similar to human performance. Second, as in humans, detection and discrimination of the stimuli are closely related. Finally, randomizing the contrast between the stimuli does not affect orientation-discrimination performance. This suggests that similar computations underlay orientation-discrimination performance in both species. In summary, our results confirm that the cat is a useful model for human orientation-discrimination performance.

Animals↗
PubMedDate not supplied

Stimulus dependence of orientation and direction sensitivity of cat LGNd relay cells without cortical inputs: a comparison with area 17 cells.

The cortical contribution to the orientation and direction sensitivity of LGNd relay cells was investigated by recording the responses of relay cells to drifting sinusoidal gratings of varying spatial frequencies, moving bars, and moving spots in cats in which the visual cortex (areas 17, 18, 19, and LS) was ablated. For comparison, the spatial-frequency dependence of orientation and direction tuning of striate cortical cells was investigated employing the same quantitative techniques used to test LGNd cells. There are no significant differences in the orientation and direction tuning to relay cells in the LGNd of normal and decorticate cats. The orientation and direction sensitivities of cortical cells are dependent on stimulus parameters in a fashion qualitatively similar to that of LGNd cells. The differences in the spatial-frequency bandwidths of LGNd cells and cortical cells may explain many of their differences in orientation and direction tuning. Although factors beyond narrowness of spatial-frequency tuning must exist to account for the much stronger orientation and direction preferences of cells in area 17 when compared to LGNd cells, the evidence suggests that the orientation and direction biases present in the afferents to the visual cortex may contribute to the orientation and direction selectivities found in cortical cells.

Animals↗
PubMed1989

Orientation and direction tuning of goldfish ganglion cells.

Orientation and direction tuning were examined in goldfish ganglion cells by drifting sinusoidal gratings across the receptive field of the cell. Each ganglion cell was first classified as X-, Y-, or W-like based on its responses to a contrast-reversal grating positioned at various spatial phases of the cell's receptive field. Sinusoidal gratings were drifted at different orientations and directions across the receptive field of the cell; spatial frequency and contrast of the grating were also varied. It was found that some X-like cells responded similarly to all orientations and directions, indicating that these cells had circular and symmetrical fields. Other X-like cells showed a preference for certain orientations at high spatial frequencies suggesting that these cells possess an elliptical center mechanism (since only the center mechanism is sensitive to high spatial frequencies). In virtually all cases, X-like cells were not directionally tuned. All but one Y-like cell displayed orientation tuning but, as with X-like cells, orientation tuning appeared only at high spatial frequencies. A substantial portion of these Y-like cells also showed a direction preference. This preference was dependent on spatial frequency but in a manner different from orientation tuning, suggesting that these two phenomena result from different mechanisms. All W-like cells possessed orientation and direction tuning, both of which depended on the spatial frequency of the stimulus. These results support past work which suggests that the center and surround components of retinal ganglion cell receptive fields are not necessarily circular or concentric, and that they may actually consist of smaller subareas.

Animals↗
PubMedDate not supplied

Heterogeneity in local distributions of orientation-selective neurons in the cat primary visual cortex.

We have employed the tetrode technique, which allows accurate discrimination of individual neuronal spike trains from multiunit recordings, in order to examine the variation of orientation selectivity among local groups of neurons. We recorded a total of 321 cells from 62 sites in area 17 of halothane-anesthetized cats; each site contained between three to ten neurons that were estimated to be less than 65 microns away from the tetrode tip. For each cell, we determined the orientation tuning in response to moving bars. Of the cells tested, 8.4% were unresponsive, 22.7% had no preferential response to any particular orientation, while 68.8% were tuned. The average difference in preferred orientation between cell pairs recorded at the same site was 10.7 deg, but the variance in preferred orientation differences differed significantly among sites. Some clusters of cells exhibited the same or nearly the same orientation preference, while others had orientation preferences that differed by as much as 90 deg. Our data demonstrate that the tuning for orientation is more heterogeneously distributed at a local level than previous studies have suggested.

Action Potentials↗
PubMedDate not supplied

Dynamics of the orientation tuning of postsynaptic potentials in the cat visual cortex.

We evaluated the dynamic aspects of the orientation tuning of the input to cat visual cortical neurons by analyzing the postsynaptic potentials (PSPs) evoked by flashing bars of light. The PSPs were recorded using in vivo whole-cell technique, and we analyzed the orientation tuning during subsequent temporal windows after stimulus onset and offset. Our results show that the amplitudes of the postsynaptic potential are reliably tuned to orientation and matching that of the spike responses only during certain temporal windows. During the first 100 ms after stimulus presentation, orientation tuning of the membrane potential underwent regular changes. Within particular intervals, orientation tuning of the input was much sharper than that estimated according to the whole response. In most cells, optimal orientation was usually stable over the whole period. In several cells which had a second hump of EPSPs in the response, this second hump was tuned to the same orientation as the first one, but always showed sharper tuning. Estimation of the integration time revealed sufficient delay between the appearance of EPSPs and spikes, to let inhibition influence spike generation. These results show that orientation selectivity of the input to cortical cells is a dynamic function, and also indicate the possibility of temporal coding in the visual system.

Animals↗
PubMedDate not supplied

Computational modeling of orientation tuning dynamics in monkey primary visual cortex.

In the primate visual pathway, orientation tuning of neurons is first observed in the primary visual cortex. The LGN cells that comprise the thalamic input to V1 are not orientation tuned, but some V1 neurons are quite selective. Two main classes of theoretical models have been offered to explain orientation selectivity: feedforward models, in which inputs from spatially aligned LGN cells are summed together by one cortical neuron; and feedback models, in which an initial weak orientation bias due to convergent LGN input is sharpened and amplified by intracortical feedback. Recent data on the dynamics of orientation tuning, obtained by a cross-correlation technique, may help to distinguish between these classes of models. To test this possibility, we simulated the measurement of orientation tuning dynamics on various receptive field models, including a simple Hubel-Wiesel type feedforward model: a linear spatiotemporal filter followed by an integrate-and-fire spike generator. The computational study reveals that simple feedforward models may account for some aspects of the experimental data but fail to explain many salient features of orientation tuning dynamics in V1 cells. A simple feedback model of interacting cells is also considered. This model is successful in explaining the appearance of Mexican-hat orientation profiles, but other features of the data continue to be unexplained.

Action Potentials↗
PubMed1990

Orientation congruency effects on the identification of disoriented shapes.

Effects of orientation on identification can be attenuated when other patterns at the same (or a similar) orientation are identified in close temporal contiguity. In Experiments 1 and 2, letters were presented simultaneously in brief masked displays. Identification accuracy was much higher when the letters had consistent orientations than when the letters had different orientations within a display. In Experiment 3, two letters were presented sequentially. Identification accuracy was higher with congruent than with incongruent orientations. The results are unexpected if one assumes that the patterns themselves are rotated until upright prior to their identification, unless pattern rotation processes can be primed, and that priming requires orientation congruence between the priming and primed stimulus. The results are expected if the orientation of a frame of reference can be adjusted to the orientation of the patterns during the identification process.

Adult↗
PubMed1986

Reinstatement of orienting behavior by d-amphetamine in rats with superior colliculus lesions.

The involvement of the nigrotectal pathway in the expression of visual orienting behavior was assessed by a combination of superior colliculus (SC) lesions and increased dopamine transmission produced by administration of d-amphetamine. Orienting behavior elicited by apparently moving or stationary light displays, its habituation, and recovery were observed. In intact animals, amphetamine injections had a small but reliable effect on the habituation of orienting behaviors. Rats with SC lesions did not orient to the lights. Amphetamine-injected rats with SC lesions did orient, and the topography of their orienting behavior, rate of habituation, and recovery of orienting with changes in the light display were comparable to those of the intact animal. These results suggest a view of SC-lesion-impaired orienting behavior as a disturbance of sensory attention and emphasize the interaction of the SC and other neural systems in processes mediating the direction of attention.

Animals↗
PubMedDate not supplied

A comparison of inhibition in orientation and spatial frequency selectivity of cat visual cortex.

Neurones in the visual cortex are highly selective for orientation and spatial frequency of visual stimuli. There is strong neurophysiological evidence that orientation selectivity is enhanced by inhibitory interconnections between columns in the cortex which have different orientation sensitivities, an idea which is supported by experiments using neuropharmacological manipulation or complex visual stimuli. It has also been proposed that selectivity for spatial frequency is mediated in part by a similar mechanism to that for orientation, although evidence for this is based on special use of visual stimuli, which hampers interpretation of the findings. We have therefore examined selectivity for both orientation and spatial frequency using a technique which allows direct inferences about inhibitory processes. Our method uses microiontophoresis of an excitatory amino acid to elevate maintained discharge of single neurones in the visual cortex. We then present visual stimuli both within and outside the range of orientations and spatial frequencies which cause a cell to respond with increased discharge. Our results show that orientations presented on either side of the responsive range usually produce clear suppression of maintained discharge. In marked contrast, spatial frequencies shown to either side of the responsive range have little or no effect on maintained activity. We conclude that there is an intracortical organization of inhibitory connections between cells tuned to different orientations but not different spatial frequencies.

Animals↗