PERCEIVED DIRECTION OF MOTION VS. DETECTION OF GLOBAL FLOW IN RANDOM DOT CINEMATOGRAMS


A.B. Sekuler, R. Sekuler*, C. Penbeci


Dept. of Psychology, U. of Toronto; *Center for Complex Systems, Brandeis U.
(based on a Poster presented at ARVO '96)

INTRODUCTION

The visual system can integrate disparate motion vectors into a unified percept of coherent motion. For example, random dot cinematograms comprising various, spatially-intermingled directions can produce clear unidirectional, global flow. This global flow is easiest to see when the cinematogram's directions span a limited range. When all possible directions are present in equal amounts, random "noisy" motions are seen instead of flow. Our research sought to answer three questions about the phenomenon of global flow.

€Question 1. What direction does global flow take? Researchers have assumed that perceived flow is in the mean of all directions present in the cinematogram (Williams & Brannan). This assumption sets strong constraints on the mechanisms responsible for global flow. Is perceived flow in the mean direction of a cinematogram, as always has been assumed?

€Question 2. How are direction and strength of global flow related to one another? The distribution of directions in a cinematogram governs the ease with which flow is seen. The same distribution presumably also governs the perceived direction of that flow. Can one framework account for both the ease of seeing global flow and the accuracy with which flow's direction tracks the overall distribution of directions?

€Question 3. What neural computations generate global flow? We evaluated two competing computational views of how the nervous system might process directional information to produce a single perceived direction of flow from a multivectoral stimulus: (1) Vector Averaging and (2) Winner-Take-All. Salzman &Newsome speculated that Vector Averaging might be responsible for the perception of global flow because a unified direction of motion is perceived; whereas the Winner-Take-All model might be responsible for motion perception under conditions in which more than one motion is perceived (e.g., transparency).




EXPERIMENT 1


Cinematograms

€300 dots; step size = 5.4 minarc; 80 msec (6 frames) display within a 6 deg circular aperture; binocular viewing from 114 cm.
€On each frame, each dot's directions were chosen randomly and independently from a uniform distribution of directions.
Signal: Distribution Width = 0, 45, 90, 135, 225, or 270 deg
Noise: Distribution Width = 360 deg.

schematic illustration of individual dot motions

Demonstrations of ONE DOT undergoing motion with a distribution width of 0 deg, 90 deg, and 180 deg. These demonstrations require Netscape 2.0

Observers' Tasks and Dependent Measures.
Direction Identification. Observers indicated perceived direction of motion by clicking on the perimeter of a circle surrounding the location of the cinematogram. Accuracy was indexed by the absolute difference between the direction response and the mean direction of the distribution.
Signal vs. Noise Discrimination. Observers used a 6-point rating scale to judge signal vs. noise, and to express confidence in judgment (Signal Definitely, Signal Probably, Signal Maybe, Noise Maybe, Noise Probably, Noise Definitely). Sensitivity (strength of flow) was indexed by 2ArcsinˆP(A) ‹ a rating scale approximation to d'.

Sequence of events on a trial


Answer 1. What direction does global flow take?
€Perceived direction is centered around a cinematogram's mean direction, but large errors are common. Accuracy decreases with distribution width.

Answer 2. How are direction and strength of global flow related?

€Within a narrow range of high sensitivity to flow, accuracy varies widely. Observers can detect flow without knowing what direction the flow took.

Answer 3. What neural computations generate global flow?
Vector averaging (VAvg). Global flow reflects a vector average of signals from directional-selective mechanisms. The direction and length of the resultant vector govern perceived direction and sensitivity, respectively. The VAvg model is supported by studies of directional adaptation and directional aftereffects (e.g., Levinson & Sekuler).
Winner-take-all (WTA). Global flow reflects the response of the single neural mechanism responding most vigorously. The center direction of that mechanism governs perceived direction; the strength of that mechanism's response governs sensitivity. In a paradigm involving the combination of visual signals and electrically stimulated neural signals in monkeys, Salzman & Newsome found support for the WTA model.

THE MODELS. In both models, directional information is represented by responses of 12 directionally-selective mechanisms. The idea of such a representation is supported by work on (i) motion metamers (Williams, Sekuler & Tweten), (ii) discrimination of global flow's direction (Watamaniuk & Sekuler), and (iii) statistical efficiency of motion perception (Watamaniuk). Mechanism n's response a cinematogram is given by the inner product between a vector containing the distribution of directions in the cinematogram, and a vector of the mechanism's sensitivity to motion in each direction.

NOISING UP THE CHANNELS. To bring the model into the domain of human observers' imperfect performance, we introduced random, Gaussian noise into the sensitivities of the 12 mechanisms. Each simulated trial used new, independent samples of Gaussian noise. We adjusted noise variance so that human and simulation accuracy matched with direction distribution width = 0. This noise level remained constant for all simulations, but the two models required different noise levels. Both models treated discriminability of signal from noise as a statistical decision based on [Rs-Rn] / sqrt[VARs+VARn], where Rs and Rn are the responses generated by a signal and noise cinematograms; and VARs and VARn are the variances of responses from signal and noise.

The next two figures illustrate the relative amount of noise introduced into directional channels for the VAvg and WTA models. Note that the VAVg model requires more noise than the WTA model to match human performance.

Illustration of relative amount of noise introduced into direction channels in the VAvg model Noised-up channels for the WTA model

RESULTS OF SIMULATIONS. Both models give respectable, if imperfect, accounts of direction accuracy. VAvg also does well in predicting observers' discrimination of signal from noise; but WTA overpredicts discriminability. This overprediction arises from the very small values of VARs produced by WTA for broad direction distributions.

Symbols in the following 4 figures represent the mean results for observers. The solid lines in each represent the predictions of the VAvg or WTA models. The first two graphs show results for direction accuracy:

Accuracy compared with VAvg predictions Accuracy compared with WTA predictions

The next two graphs show results for sensitivity:

Sensy compared with VAvg predictions Sensy compared with WTA predictions

Take Home Message from Experiment 1: The Vector Averaging model provides a good fit for both accuracy and sensitivity. The Winner- Take-All model does not predict both sets of results. The visual system appears to rely on a neural computation like that of Vector Averaging when integrating disparate direction signals.

Question for Experiment 2: Does the visual system always rely on Vector Averaging?



EXPERIMENT 2

Continuity of directions
€In some everyday conditions the directions present within a local region do not form a continuous set. (e.g., perceptual transparency). In these conditions, WTA is more plausible; VAvg discards directional information needed to generate more than one perceptual flow (Salzman & Newsome).
€The visual system often can "fill in" information missing from the retinal image (Ramachandran). Such filling in typically operates across spatial gaps.
€Can the visual system fill in missing directions of motion from some otherwise contiguous set of motions? Or do all gaps lead to percepts of transparency?
€Gap stimuli give the WTA model a strong test, particularly when possible directions of flow are highly similar. Adjacent directions in Salzman & Newsome's study were 45 degrees apart, which may not be a real challenge for the model.
€Gap stimuli also test the degree of flexibility for the VAvg model. If VAvg fails, does visual system shift to WTA?
€We examined human observers' responses to cinematograms with directional gaps, and also compared VAvg and WTA models' responses to these stimuli.

Cinematograms and Procedure
Stimuli and tasks were as in Experiment 1, except:
€Observers discriminated: (i) a signal distribution (with a gap centered around the mean direction) from the same distribution width with no gap (noise). The total number of dots was equal across gap and no-gap stimuli.
€Within each session, distribution width was constant (90, 135, or 180 deg), but gap size varied (from 0 to 135 deg, in 15 deg steps).

Results: Identification of Direction
€Errors in direction identification increased with gap width; the rate and extent of this increase varied among observers.
€WTA provided a better fit than VAvg for all observers with Distribution Width = 90, for ABS with DW = 135, and for MBS with DW = 180.
€Vavg provided a better fit than WTA for CP with Distribution Widths = 135 and 180, although in these cases the fits declined with increasing gap width.
Neither model consistently performed well.

These figures show the accuracy (relative to the mean of the distribution) for perceived direction for Gap stimuli with Distribution widths of 90, 135, and 180 deg. In all figures, the lower line shows predictions of the VAvg model, and the upper line shows predictions of the WTA model.
Accuracy for distribution width=90 Accuracy for distribution width=135 Accuracy for distribution width=180

Results: Discrimination of Gap vs. No-gap
€Discrimination of gap from non-gap cinematograms improved with increasing gap width, but the rate of improvement varied considerably among observers.
€Regardless of the cinematogram's overall width of direction distribution, a constant gap width produced a constant level of discriminability.
Neither model predicted the obtained pattern of discrimination results.

The next three figures show sensitivity as a function of gap width for observers ABS, CP, and MBS.
ABS CP MBS

The next two figures show the predictions of the VAvg and WTA models. Note that the scales have been altered significantly relative to observers.
VAvg WTA

The Bottom Line for Experiment 2: Neither the Vector Averaging nor the Winner-Take-All models consistently predict the accuracy and sensitity of observers. The introduction of a gap -- and of perceived transparency -- causes the visual system to use a neural computation other than Vector Averaging. However, contrary to previous predictions, the visual system does not use a Winner-Take-All strategy either.



GENERAL DISCUSSION


Phenomenology & Individual differences
€Subjects usually were certain about their direction identifications, no matter how inaccurate the identifications were.
€Gap stimuli sometimes took on an unstable appearance, switching between global flow and perceptual transparency.
€Continuous distributions of direction produced good agreement among observers, but discontinuous distributions produced considerable divergence among observers. Individual differences may derive from differences in categorizing stimuli that potentially are bistable.

Relationship between WTA and VAvg

The models we implemented are the simplest representatives of their respective classes. Each had few parameters that needed to be estimated, and both were silent about spatial and temporal variables known to affect perception of motion (Williams, Phillips & Sekuler; Nawrot & Sekuler; Braddick; McKee & Watamaniuk). The two classes of models are not mutually exclusive, but lie along one continuum. In a variant VAvg, non-linear averaging would lend extra weight to mechanisms that gave the strongest responses. In the extreme, the non-linearity would silence all but the mechanism responding most strongly (WTA).

Flexibility
The visual system may use computations that resemble WTA as well as computations that resemble VAvg, with the two giving rise to different percepts: global flow and transparency. The two modes of processing may represent two different stages of processing, and those stages may interact. Such an interaction would be analogous to Leopold & Logothetis' account of binocular rivalry and other multistable perceptual states. It would also be consistent with the bistable responses evoked by our gap cinematograms.

Ideal Behaviour
Sensitivity results from Experiment 2 suggest that observers can adapt their behaviour ideally. Regardless of distribution range, gap width was the primary predictor of sensitivity. This pattern is expected from an ideal observer, although neither the VAvg nor WTA models produce this result. Further research is required to determine the extent to which observers can control the strategies with which they perceive motion, or attend to specific aspect of a stimulus.



References

Braddick OJ (1993) Segmentation versus integration in visual motion processing. Trends in Neuroscience 16 263-268.

Leopold DA & Logothetis NK (1996) Activity changes in early visual cortex reflect monkeys' percepts during binocular rivalry. Nature 379 549-553.

Levinson E & Sekuler R (1980) A two-dimensional analysis of direction-specific adaptation. Vision Research 20 103-107.

McKee SP & Watamaniuk SNJ (1994) The psychophysics of motion perception. In AT Smith & RJ Snowden (eds.) Visual detection of motion. Academic Press. Pp. 85-114.

Nawrot M & Sekuler R (1990) Assimilation and contrast in motion perception: explorations in cooperativity. Vision Research 30 1439-1451.

Ramachandran VS (1993) Filling in gaps in perception. Part II. Scotomas and phantom limbs. Current Directions in Psychological Science 2 56-65.

Salzman CD & Newsome WT (1994) Neural mechanisms for forming a perceptual decision. Science 264 231-237.

Snowden RJ, Treue S, Erickson RG & Andersen RA (1991). The responses of Area MT and V1 neurons to transparent motion. Journal of Neuroscience 11 2768-2785.

Watson AB & Robson JG (1981) Discrimination at threshold: Labelled detectors in human vision. Vision Research 21 1115-1122.

Watamaniuk SNJ, Sekuler R & Williams DW (1989) Direction perception in complex visual displays: The integration of direction information. Vision Research 29 47-59

Williams DW, Tweten S & Sekuler R (1991) Using metamers to explore motion perception. Vision Research 31 275-286.

Williams DW & Brannan JR (1994) Spatial integration of local motion. In AT Smith& RJ Snowden (eds.) Visual detection of motion. Academic Press. Pp. 291-303.



SUPPORTED by the Natural Sciences and Engineering Research Council of Canada, the James S. McDonnell Foundation, and the M. R. Bauer Foundation.



QUESTIONS/COMMENTS TO

Allison Sekuler:

sekuler@psych.utoronto.ca

or

Robert Sekuler:

sekuler@volen.ccs.brandeis.edu