Horizontal-vertical filters in early vision predict anomalous line-orientation identification frequencies.

A characteristic of early visual processing is a reduction in the effective number of filter mechanisms acting in parallel over the visual field. In the detection of a line target differing in orientation from a background of lines, performance with brief displays appears to be determined by just two classes of orientation-sensitive filter, with preferred orientations close to the vertical and horizontal. An orientation signal represented as a linear combination of responses from such filters is shown to provide a quantitative prediction of the probability density function for identifying the perceived orientation of a target line. This prediction was confirmed in an orientation-matching experiment, which showed that the precision of orientation estimates was worst near the vertical and horizontal and best at about 30 degrees each side of the vertical, a result that contrasts with the classical oblique effect in vision, when scrutiny of the image is allowed. A comparison of predicted and observed frequency distributions showed that the hypothesized orientation signal was formed as an opponent combination and horizontal and vertical filter responses.     

Multiple groups of orientation-selective visual mechanisms underlying rapid orientated-line detection.

Visual search for an edge or line element differing in orientation from a background of other edge or line elements can be performed rapidly and effortlessly. In this study, based on psychophysical measurements with ten human observers, threshold values of the angle between a target and background line elements were obtained as functions of background-element orientation, in brief masked displays. A repeated-loess analysis of the threshold functions suggested the existence of several groups of orientation-selective mechanisms contributing to rapid orientated-line detection; specifically, coarse, intermediate and fine mechanisms with preferred orientations spaced at angles of approximately 90 degrees, 35 degrees, and 10 degrees-25 degrees, respectively. The preferred orientations of coarse and some intermediate mechanisms coincided with the vertical or horizontal of the frontoparallel plane, but the preferred orientations of fine mechanisms varied randomly from observer to observer, possibly reflecting individual variations in neuronal sampling characteristics.     

Perceptual segregation and retinal position

This study is concerned with the dependence of perceptual segregation performance on the retinal position at which the performance is evaluated. Segregation performance consisted in detecting a target texture composed of line elements of constant length and orientation embedded in a background texture. The background texture was made up of the same line elements as the target texture but the background line elements were set at 90 deg to the target elements. Results showed that, at least for 40- and 50-ms presentation times, this task could be much more effectively completed outside the fovea centralis than within this area. These findings indicate that extrafoveal areas of the retina may make a significant and previously underestimated contribution to perceptual segregation.     

A space-variant filter model of texture segregation: Parameter adjustment guided by psychophysical data

 This article presents a space-variant version of a standard spatial filter model of texture segregation of the “back-pocket” type (i.e., two filter layers with an intermediate pointwise nonlinearity). The model was tested with psychophysical data from experiments with line textures in which target lines differed in orientation from background lines. The textures were presented briefly and then masked. Segregation performance was evaluated along the horizontal meridian up to retinal eccentricities of about 10 deg. Data are reported from two experiments with different line densities (Kehrer 1989) and two experiments with different orientation contrasts between target lines and background lines (Kehrer 1990). Segregation performance proved to depend strongly on these texture variations, and it peaked several degrees from fixation in all cases. The filter model provided satisfactory predictions of experimental data when model parameters were adjusted appropriately. It is concluded (1) that filter models defined in strictly spatial terms (i.e., without temporal properties) offer a sufficient framework to account for the psychophysical data and (2) that the particular course of the performance curve (i.e., the performance peak outside the central region) must be attributed to the characteristics of second-layer filters. Received: 28 June 2001 / Accepted in revised form: 10 October 2002 / Published online: 13 February 2003 Correspondence to: L. Kehrer (e-mail: lothar.kehrer@uni-bielefeld.de) Acknowledgements. This work was supported by Grant Ke 388/3-2 from the Deutsche Forschungsgemeinschaft (DFG). We wish to thank Jonathan Harrow for improving the English text and an anonymous reviewer for many constructive comments.     

Two spatio-temporal filters in human vision

1. We have studied visual detection of a circular target moving across a spatially and/or temporally modulated background. Illumination, I _t , for threshold detection of the target has been measured as a function of background modulation frequency and changes in I _t associated with background modulation provide a means of determining the frequency response characteristics of visual channels. 2. Temporal frequency responses obtained with temporally modulated, spatially uniform backgrounds have pass-band characteristics and the temporal frequency for peak response increases with increase in mean background illumination. These temporal frequency responses resemble those of the de Lange (1954) filter, but the latter incorporates the incremental thresholds for steady backgrounds. 3. The amplitude of this temporal response saturates at low (40%) background modulation, decreases to zero as the target velocity falls to zero, and is maximum for a circular target of diameter 2°. 4. The spatial characteristics of this temporal filter were measured with a background field consisting of alternate steady and flickering bars. The resulting spatial frequency curve peaks at 1 cycle deg^-1 for all background illuminations and is independent of the background grating orientation. This spatial response differs significantly from the IMG spatial functions observed with a background grating (Barbur and Ruddock, 1980). 5. The spatial and temporal responses reviewed above exhibit similar parametric variations and we therefore associate them with a single spatiotemporal filter, ST2. 6. A second temporal response, with low-pass frequency characteristics, was observed with a background field consisting of two matched gratings, presented in spatial and temporal antiphase. This response has parametric properties similar to those of the IMG spatial response described previously by Barbur and Ruddock (1980), thus we associated the two sets of data with a single spatio-temporal filter, ST1. 7. We show that the ST2 responses can be obtained by combining ST1 responses, and we present a network incorporating the two filters. 8. We review other psychophysical studies which imply the activity of two spatio-temporal filters with properties of the kind revealed in our studies. We argue that filter ST1 has properties equivalent to those of X-type and filter ST2 has properties equivalent to those of Y-type electrophysiological mechanisms.     

Local spatial scale for three-dot alignment acuity

Three-dot alignment discrimination thresholds were determined for blobs with Gaussian spatial and temporal contrast envelopes. The stimuli were presented at detection threshold luminance contrast. Thresholds were determined as a function of the blur parameter of the stimuli. This was done for a range of eccentricities in the visual field (from 45 degrees nasal to 65 degrees temporal). The thresholds were corrected for variations of the stimulus extent with the blur parameter. The results were used to estimate the local spatial scale for three-dot alignment acuity. This was done by a method recently introduced by Watson (1987). It was found that the local spatial scale for three-dot alignment acuity is approximately linearly proportional to eccentricity.     

Collinear effects on 3-Gabor alignment as a function of spacing, orientation and detectability.

Our ability to align three Gabor patches depends upon their internal carrier orientation; we are better at aligning vertical or horizontal patches than oblique patches (Keeble and Hess, 1998). However, the tuning of alignment to patch orientation has not studied in detail. We measured the alignment of a vertical target with reference patches varying in orientation and found it tuned to vertical (collinear) patches at centre-to-centre separation of three carrier periods, with a steep increase for oblique references and slight downturn for horizontal (orthogonal) references. Next, we increased separation between the patches, testing collinear, side-by-side, orthogonal and oblique configurations. Surprisingly, we found that the tuning for collinear patches was preserved. All ten observers tested had lower alignment thresholds for collinear patches. This effect extended to an inter-patch separation of 10 carrier periods (20 envelope standard deviations). Additionally, we measured contrast detection thresholds for the reference patches using the same stimuli. The collinear facilitation of alignment was even greater than the collinear facilitation of detection.     

The central performance drop in texture segmentation: a simulation based on a spatial filter model

 This article presents a computational model of early visual information processing that attempts to account for the central performance drop (CPD) in texture segmentation. CPD is the finding that detection performance on short stimulus displays of line textures using orientation differences to set off the target is not maximal at the foveal center but in parafoveal areas. A comparison between a simulation and psychophysical experimental data supported the assumption that the CPD may be explained by properties of spatial frequency channels whose band-pass filter characteristics are not constant over the retina but differ with eccentricity in a defined manner. The model provided satisfactory predictions of experimental data based on densely or widely spaced line elements in texture fields. It is concluded that preattentive texture analysis might be performed by a relatively small number of simple spatial filters. Received: 14 November 1996 / Accepted in revised form: 3 June 1997     

Do alignment thresholds define a critical boundary in long-range detection facilitation with co-linear lines?

Thresholds for line contrast detection (experiment 2) were measured with a two-alternative temporal forced-choice procedure as a function of the spatial position of a vertical target line with regard to two co-linear context lines. The different spatial positions of the target line corresponded to values near the position discrimination threshold (experiment 1) reflecting the just detectable lateral offset, or non-co-linearity, between the context lines which were vertically separated by about 100 minutes of visual arc. Target and context lines were vertically separated by about 30 minutes of arc, had equal contrast polarity in one case, and opposite contrast polarity in the other. Strong line contrast detection facilitation is found at perceptually co-linear target locations. This facilitation decreases noticeably at a horizontal target offset that corresponds to the alignment threshold measured with the context lines. The effects are independent of the relative contrast polarity of target and context and, as shown in a third experiment, also independent of both the relative length or number of lines, and the magnitude of their absolute co-axial separation. This independence seems to hold, provided individual line length and co-axial distance between lines are larger than what appears to be the lower limit of the long-range spatial domain for orientation or contour integration (i.e. 20 minutes of arc), as determined by previous studies. The findings reported here suggest that alignment thresholds are likely to define a critical lateral boundary in long-range detection facilitation with co-linear lines. They support models of contour integration based on interactions between neural mechanisms that integrate local signals of contrast, orientation, and relative position or end-to-end alignment. Such mechanisms may help to explain the formation of representations of virtual contours and object contours in human perception.     

Sound localization by the bottlenose porpoise Tursiops truncatus.

1. Sound localization was measured behaviourally for the Atlantic bottlenose porpoise (Tursiops truncatus) using a wide range of pure tone pulses as well as clicks simulating the species echolocation click. 2. Measurements of the minimum audible angle (MAA) on the horizontal plane give localization discrimination thresholds of between 2 and 3 degrees for sounds from 20 to 90 kHz and thresholds from 2-8 to 4 degrees at 6, 10 and 100 kHz. With the azimuth of the animal changed relative to the speakers the MAAs were 1-3-1-5 degrees at an azimuth of 15 degrees and about 5 degrees for an azimuth of 30 degrees. 3. MAAs to clicks were 0-7-0-8 degrees. 4. The animal was able to do almost as well in determining the position of vertical sound sources as it could for horizontal localization. 5. The data indicate that at low frequencies the animal may have been localizing by using the region around the external auditory meatus as a detector, but at frequencies about 20 kHz it is likely that the animal was detecting sounds through the lateral sides of the lower jaw. 6. Above 20 kHz, it is likely that the animal was localizing using binaural intensity cues. 7. Our data support evidence that the lower jaw is an important channel for sound detection in Tursiops.     

Detection of a gabor patch superimposed on an illusory contour

Interactions between visual stimuli have been found to be specific to the spatial frequency, orientation and phase of the interacting stimuli. We asked if there are any interactions between luminance-defined Gabor patches and Kanizsa-type illusory contours. In psychophysical experiments we studied whether induction of a vertical illusory line affects detection thresholds for a Gabor patch superimposed on this line and whether these effects depend on the orientation, spatial frequency and phase of the Gabor elements. Employing a 2AFC method with a staircase procedure we measured contrast detection thresholds and varied the orientation, spatial frequency and phase of the test Gabor patch and the separation between the two pacmen in four experimental series. The results show that in a situation where the two inducers generate perception of an illusory line, the contrast detection of the Gabor patch is facilitated relative to a control condition where the rotated pacmen do not induce illusory contours. This facilitation was more pronounced for test Gabor signals that were collinear to the illusory line, but the observer's performance was not altered by changes in the spatial frequency or phase of the Gabor stimuli. With increasing spatial separation of the two pacmen (and, consequently, with a decreasing support ratio), the difference between performance in the test and control conditions diminished. From the data obtained we cannot infer that we have measured some neural interactions between Gabor patches and Kanizsa-type illusory contours, and nor can we draw a unique conclusion about what causes the facilitation of detection of the test Gabor patch in the experimental situation that allows induction of the illusory line. We discuss possible mechanisms of the facilitation, such as contextual influences or a reduction of uncertainty about spatial location of the test Gabor patch.     

Short-range limitation on detection of feature differences

We studied the ability of observers to detect the presence of a clearly visible line segment against a background of line segments of different orientation. As we increase the number (density) of these background lines, we find that detectability does not behave monotonically. Adding a small number of background lines decreases detectability but if adjacent line segments are permitted to fall in close range, a further increase of background lines improves performance which eventually reaches a constant level. This suggests that detection of feature differences involves a short-range process. The range of this process is about two degrees or twice the length of the line segments used. Thus texture-gradients between different elements are only formed if the distance between these elements is not much larger than the average element size.     

Time course analysis of baroreflex sensitivity during postural stress

Postural stress requires immediate autonomic nervous action to maintain blood pressure. We determined time-domain cardiac baroreflex sensitivity (BRS) and time delay (tau) between systolic blood pressure and interbeat interval variations during stepwise changes in the angle of vertical body axis (alpha). The assumption was that with increasing postural stress, BRS becomes attenuated, accompanied by a shift in tau toward higher values. In 10 healthy young volunteers, alpha included 20 degrees head-down tilt (-20 degrees), supine (0 degree), 30 and 70 degrees head-up tilt (30 degrees, 70 degrees), and free standing (90 degrees). Noninvasive blood pressures were analyzed over 6-min periods before and after each change in alpha. The BRS was determined by frequency-domain analysis and with xBRS, a cross-correlation time-domain method. On average, between 28 (-20 degrees) to 45 (90 degrees) xBRS estimates per minute became available. Following a change in alpha, xBRS reached a different mean level in the first minute in 78% of the cases and in 93% after 6 min. With increasing alpha, BRS decreased: BRS = -10.1.sin(alpha) + 18.7 (r(2) = 0.99) with tight correlation between xBRS and cross-spectral gain (r(2) approximately 0.97). Delay tau shifted toward higher values. In conclusion, in healthy subjects the sensitivity of the cardiac baroreflex obtained from time domain decreases linearly with sin(alpha), and the start of baroreflex adaptation to a physiological perturbation like postural stress occurs rapidly. The decreases of BRS and reduction of short tau may be the result of reduced vagal activity with increasing alpha.     

Orientation dependence of displacements by a single one-headed myosin relative to the actin filament.

Displacements of single one-headed myosin molecules in a sparse myosin-rod cofilament were measured from bead displacements at various angles relative to an actin filament by dual optical trapping nanometry. The sparse myosin-rod cofilaments, 5-8 micron long, were synthesized by slowly mixing one-headed myosin prepared by papain digestion with myosin rods at molar ratios of 1:400 to 1:1500, so that one to four one-headed myosin molecules were on average scattered along the cofilament. The bead displacement was approximately 10 nm at low loads ( approximately 0.5 pN) and at angles of 5-10 degrees between the actin and myosin filaments (near physiologically correct orientation). The bead displacement decreased with an increase in the angle. The bead displacement at nearly 90 degrees was approximately 0 nm. When the angle was increased to approximately 150 degrees-170 degrees, the bead displacements increased to 5 nm. A native two-headed myosin showed similar size and orientation dependence of bead displacements as a one-headed myosin.     

Orientation selectivity and visual acuity in schoolchildren and adults

We continued our study of the mechanisms of visual acuity (VA) in ontogenesis. We measured the VA and sensitivity to lines orientation and determined the minimal length of lines for discrimination of the vertical and the horizontal ones in subjects aged 7–25 years. It was found that the thresholds of discrimination of vertical and horizontal lines in subjects with normal VA decreased with age up to 9–10 years and then remained constant, while the orientation selectivity was improving up to the age of 14–16 years. The average VA almost did not depend on age. Individual thresholds of line lengths and orientation discrimination correlated with the VA of subjects.     

Echo thresholds measured in the vertical and horizontal planes

The results of studying the precedence effect in the case where the direct and delayed (reflected) signals are located in the vertical and horizontal planes are considered. Loudspeakers emitting direct and reflected sounds were placed 45 deg to the left and right of the median line of the subject’s head in the horizontal plane and in front of and above the subject’s head, i.e., with 0 and 90 deg of elevation relative to the eye level, in the vertical plane. It has been shown that the time limits of the precedence effect of short (5-ms) signals are similar in the horizontal and vertical planes. For signals more than 10 ms in duration, the values of echo thresholds were higher in the vertical plane and significantly differed (p < 0.05) from the thresholds in the horizontal plane.     

Feature matching and segmentation in motion perception

We examined the role of feature matching in motion perception. The stimulus sequence was constructed from a vertical, 1 cycle deg-1 sinusoidal grating divided into horizontal strips of equal height, where alternate strips moved leftward and rightward. The initial relative phase of adjacent strips was either 0 degree (aligned) or 90 degrees (non-aligned) and the motion was sampled at 90 degrees phase steps. A blank interstimulus interval (ISI) of 0-117 ms was introduced between each 33 ms presentation of the stimulus frames. The observers had to identify the direction of motion of the central strip. Motion was perceived correctly at short ISIs, but at longer ISIs performance was much better for the non-aligned sequence than the aligned sequence. This difference in performance may reflect a role for feature correspondence and grouping of features in motion perception at longer ISIs. In the aligned sequence half the frames consisted of a single coherent vertical grating, while the interleaved frames contained short strips. We argue that to achieve feature matching over time, the long edge and bar features must be broken up perceptually (segmented) into shorter elements before these short segments can appear to move in opposite directions. This idea correctly predicted that overlaying narrow, stationary, black horizontal lines at the junctions of the grating strips would improve performance in the aligned condition. The results support the view that, in addition to motion energy, feature analysis and feature tracking play an important role in motion perception.     

Slant-tilt: The visual encoding of surface orientation

A specific form for the internal representation of local surface orientation is proposed, which is similar to Gibson's (1950) “amount and direction of slant”. Slant amount is usually quantifed by the angle σ between the surface normal and the line of sight (0°≦σ≦90°). Slant direction corresponds to the direction of the gradient of distance from the viewer to the surface, and may be defined by the image direction τ to which the surface normal would project (0°≦τ≦360°). Since the direction of slant is specified by the tilt of the projected surface normal, it is referred to as surface tilt (Stevens, 1979; Marr, 1982). The two degrees of freedom of orientation are therefore quantified by slant, an angle measured perpendicular to the image plane, and tilt, an angle measured in the image plane. The slanttilt form provides several computational advantages relative to some other proposals and is consistent with various psychological phenomena. Slant might be encoded by various means, e.g. by the cosine of the angle, by the tangent, or linearly by the angle itself. Experimental results are reported that suggest that slant is encoded by an internal parameter that varies linearly with slant angle, with resolution of roughly one part in 100. Thus we propose that surface orientation is encoded in human vision by two quantities, one varying linearly with slant angle, the other varying linearly with tilt angle.     

Anisotropy in the chromatic channel: a horizontal-vertical effect

We compared the chromatic contrast thresholds of drifting (2Hz) red-green sine-wave gratings of horizontal, vertical, and two oblique orientations at three spatial frequencies (2, 4, 8 cpd). Luminance contrast thresholds for yellow-black gratings were also obtained. The classic oblique effect was found for high spatial frequency luminance and chromatic stimuli. For chromatic thresholds, a significant difference was found between the horizontal and vertical thresholds of all observers. One observer was retested with her head tilted 45 deg and demonstrated that the anisotropy was specific to retinal coordinates. These results give evidence for orientation selectivity in the chromatic channel which is at least partially independent of that in the luminance channel. We estimated the degree of lateral chromatic aberration in our observers' eyes and discuss the possible contribution of this aberration to the horizontal-vertical difference in the chromatic channel.