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Orientation-selective adaptation and tilt after-effect from invisible patterns

Abstract

Exposure to visual patterns of high contrast (for example, gratings formed by alternating white and black bars) creates after-effects in perception. We become temporarily insensitive to faint test patterns that resemble the pre-exposed pattern (such as gratings of the same orientation), and we require more contrast to detect them1. Moreover, if the test pattern is slightly tilted relative to the pre-exposed one, this tilt may be perceptually exaggerated: we experience a tilt after-effect2,3. Here we show that these visual after-effects occur even if the pre-exposed grating is too fine to be perceptually resolved. After looking at a very fine grating, so high in spatial frequency that it was perceptually indistinguishable from a uniform field, observers required more contrast to detect a test grating presented at the same orientation than one presented at the orthogonal orientation. They also experienced a tilt after-effect that depended on the relation of the test pattern's tilt to the unseen orientation of the pre-exposed pattern. Because these after-effects are due to changes in orientation-sensitive mechanisms in visual cortex4,5,6, our observations imply that extremely fine details, even those too fine to be seen, can penetrate the visual system as far as the cortex, where they are represented neurally without conscious awareness.

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Figure 1: Orientation-selective adaptation experiment.
Figure 2: Tilt after-effect experiment.

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Acknowledgements

We thank H. Smallman for his help during the experiment. This work is supported by grants from NIH and Alfred Sloan Foundation.

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Correspondence to Sheng He.

Supplementary information

1. Stimuli parameters:

The grating stimuli were presented through a laser interferometer built at UCSD, which was described in more detail in (He, S and MacLeod, D. I. A. (1996) Local luminance nonlinearity and receptor aliasing in the detection of high frequency gratings. Journal of the Optical Society of America, A. 13, 1139-1151). The interferometer allows us to introduce any level of spatial contrast to the field without altering its mean luminance. In this study, the stimuli were presented in a circular filed of 3 degrees diameter, with the mean luminance remained constant at 1000 trolands.

2. Forced-choice contrast threshold measurement procedure and results

The threshold contrasts for vertical and horizontal testing gratings after adaptation were measured with a 2AFC staircase procedure. In each session, subjects adapted to only one orientation (either horizontal or vertical), but measured contrast thresholds for both horizontal and vertical testing orientations. For each adapting spatial frequency (e.g., 65 cpd), two sessions were required to obtain 4 data points (see Figure A below).

Figure A (GIF 49.4 KB)

The staircase procedure was described in the paper itself (see figure 1a in the paper). Initial adaptation duration was 60 sec without interruption. Afterwards, a test stimulus was presented once every 6 s This test grating could be either horizontal or vertical in orientation, and could be either presented in the first or the second temporal intervals marked by audio clicks. Subjects task was to report in which interval a stimulus was presented. Based on the subjects response, the contrast in the next trial of the same orientation was determined. The two staircases (both of which were 3 up,1 down) for the horizontal and vertical test gratings were intermixed but independent of each other. 100 trials were run for each test orientation. The results (% correct at each contrast level) were fitted with the integral of a normal Gaussian function of (contrast/sigma)^n, where sigma is given as one parameter and n as the second, with the 50% point pinned at contrast = 0. The contrast level that yielded 84% correct responses was taken as the threshold, and the standard error of the threshold estimate was based on the marginal distribution of the likelihood of the data as a function of the fitted curves 84% point.

To illustrate how the threshold elevation was derived, SHs data at adapting frequency of 65 cpd are plotted in Figure AAfter adapting to a 65 cpd horizontal grating, SH measured his contrast threshold with a horizontal and a vertical grating, both at 45 cpd, using the 2AFC method described above (square symbols in Figure A). In a different session, similarly 2AFC experiments were run with a vertical 65 cpd adapting grating, generating another two data points (circles in Figure A). These two sessions generated 4 data points (2 adapting orientations x 2 testing orientations), as plotted in Figure A. Although this particular observer had a higher contrast threshold for vertical orientation in general, the interaction term in this 2x2 plot represents the orientation-specific adaptation effect. The difference of log contrast thresholds of the horizontal testing grating between horizontal and vertical adapting gratings, plus the difference of the log contrast thresholds of the vertical testing grating between vertical and horizontal adapting gratings was used as the measure of orientation-specific adaptation effect. Four independent sessions of threshold measurements were made for each adapting orientation, resulting in 4 threshold elevation values. The mean and standard error of these 4 values were plotted in Figure 1b in the paper.

3. Tilt-Aftereffect measurement procedure and results:

The stimuli parameters (luminance and filed size) were the same in this experiment as in the threshold measurement experiment. The experiment was blocked with a constant adapting orientation (15 degrees away from horizontal, either clockwise or counterclockwise) in each block. However, because the grating was not resolvable and we automated the sequence of testing blocks of different adapting orientations, the adapting orientation varied from block to block without the observers knowledge. During each block of trials, a staircase procedure (1 up and 1 down) was used to search for the testing orientation that resulted in subjective horizontal of the testing orientation. There were 50 trials in each block. The observer viewed the adapting grating (e.g., 65 cpd, 15 degree clockwise from horizontal) for 60 s at the beginning of the block, then the testing grating (at 48 cpd) was presented for 250 ms every 5.75 s, at a random orientation between + and 30 deg from horizontal. The subject pressed one of two mouse buttons to indicate whether the test grating was tilted clockwise or counterclockwise from horizontal. The staircase procedure selected the testing orientation for the next trial based on this response. In other words, subjects response drove the staircase to an eventual orientation that was subjectively horizontal. The frequency of "clockwise judgments was fit, using a maximum likelihood criterion, to a cumulative Gaussian function of physical tilt, with the 50% point and the standard deviation of the Gaussian as free parameters. The 50% point of the best fitting Gaussian was taken as the subjective horizontal, and the standard error of the estimate was based on the marginal distribution of the likelihood of the data as a function of the fitted curves 50% point.

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He, S., MacLeod, D. Orientation-selective adaptation and tilt after-effect from invisible patterns. Nature 411, 473–476 (2001). https://doi.org/10.1038/35078072

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