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Motion Sensitivity Visual Motion: Visual motion, at the most basic level, is a change in the position of a luminance pattern over time. The velocity is expressed in degrees of visual angle/second. Humans are very sensitive at detecting motion in visual images. The smallest motion that humans have been reported to detect is about 10 arc seconds, which is similar to the best threshold for vernier acuity. Motion detection is therefore like other hyperacuities, the threshold for the smallest detectable shift is much less than the size of a single cone. Motion is a correlation across space and time: In order for us to see visual motion, our brains have to compare the activity of individual receptors across space and time. One cone cannot see motion! One cone can only see light increasing or decreasing at it’s location. Motion comes from a comparison of two or more detectors at different locations, with activity that is a little delayed from one to the next. See the demo GratingThroughHoles.avi to illustrate what individual cones see when a grating drifts across the retina. Apparent Motion: Although objects in the world move continuously across our visual field, continuous real motion is not required for the perception of motion. When two stimuli appear with the correct separation in time and space, our brains link them together into “Apparent Motion” and we see them as a single moving object. The phenomenon is variously called Apparent Motion, Sampled Motion, Phi-phenomenon and/or Beta-phenomenon. The last two terms were coined by Max Wertheimer in his 1912 studies of motion perception. See demo of Phi phenomenon at http://www1.psych.purdue.edu/Magniphi/SimpliPhi.html SpatioTemporal Contrast Sensitivity and Motion When the moving object is a sine wave grating, we can make a meaningful link between spatial frequency, temporal frequency, and motion. For example, if a 1 cycle per degree grating drifts at 10 degrees per second, then the luminance at each point in the display is oscillating sinusoidally at 10 Hz. If a 10 cycle per degree grating drifts at 1 degree per second, that will also produce 10 Hz flicker at each point. See the demo movies driftgrat4Hz32cpi.avi driftgrat1Hz8cpi.avi and driftgrat1Hz2cpi.avi. The first two have the same speed (8 seconds for a bar to cross the picture), but the last two have the same temporal frequency (1 Hz flicker at each point in the image). Detection of drifting gratings follows the same pattern as we have seen for flickering gratings, once converted to temporal frequency. Current thinking is that flicker is actually detected by motion mechanisms in most cases. The figure at right shows Contrast Sensitivity for detecting drifting gratings of various spatial frequencies and drift speeds. Each motion stimulus is plotted as a function of its equivalent RESEARCH NOTE temporal frequency. For example, SPATIAL a 1 cpd grating moving at 10 PHASE SENSITIVITY FOR SINUSOIDAL GRATING TARGETS deg/sec makes a 10 Hz flicker at each point. A grating that is 0.1 GERALD WESTHEIMER cpd that moves at 100 degrees per University of California. Berkeley, CA 94770, U.S.A. Department of Physiology-Anatomy. second also makes a 10 Hz (Rrceiced 30 Nouember 19771 flicker. Except for the slowest motion (1 Abstract-Because the full specification of a sinusoidal grating target must include phase as well as deg/sec) the curves all was performed to determine the phase sensitivity for sinusoidal gratings amplitude, an experiment in central vision for normal human observers. Phase displacement thresholds for sinusoidal gratings superimpose, showing that ranging in spatial frequency from 3 to 25 cycles/degree are a linear function of spatial frequency (1’ of frequency phase angle per c’deg). The just-detectable lateral distances are constant and equal to those temporal is the for a single line. important variable for detecting the moving gratings at contrast threshold. (Data from Burr & modulation thresholds for sinusoidal grating targets This problem was approached by studying the senRoss 1982) have been investigated thoroughly. but the full specilisitivity of two normal human observers to suddenly cation of sinusoids of given frequencies requires information as to both amplitude and phase. The nature Minimum Motion Threshold: of phase sensitivity has important consequences for thresholds are a mechanmeasure theMinimum identificationmotion of the underlying detection imposed phase changes in suprathreshold sinusoidal grating patterns. The subject3 task was to identify the direction of the phase change. Seven stimuli were of the smallest displacement of aand pattern presented in random order: no shift; shifts that of one, in twothe or range three times a basic moduleand either the typically of arc seconds dotonot a subject can just identify. Values are depend much on spatial frequency above 5 cycles per degree. This figure from Westheimer (1978, Vision Research) shows minimum detectable motion for a high contrast grating pattern that jumps suddenly from one position to another. Subjects had to say if the grating jumped left or right, and the smallest displacement that produced 75% correct responses was taken as threshold. All gratings from 5 to 30 cycles per degree showed the same threshold of about 10 arc seconds. Thresholds for a single thin line gave similar results. The upper figure plots these results as “phase angle” changes, where one grating cycle = 360° of phase. The lower figure plots the same data as displacement in visual angle. Threshold for seeing the smallest motion is the same for all spatial 0 IO 20 SPATIAL FREQUENCY frequencies tested(CYCLES/ here.DEGREE) SPATIAL FREQUENCY KYCLh’DEGREEl 7 Fig. 1. (Upper) ~~~~i~~~~~poinrs are the thresholds for detection of sudden change in spatia1 phase of sinusoidal gratings, 0.5’ high. contrast 50”:. centrally fixated. for two subjects with binocular observation. Results expressed in degrees of phase angle for each grating. Vertical brackets: f 1 standard error of the mean. Srruighr lines show the mean (solid line) kl standard error (interrupted lines) for the threshold of detection of direction of a lateral displacement of a single line target. graphed as the phase angle of its Fourier components. (Lower) Same data, but displacement thresholds are graphed in the measure of linear distance rather than phase angle. 1073 Motion Aftereffect Our motion perception shows adaptation effects similar to color, orientation, brightness and even faces. Stare at a strong stimulus for a period of time, it will seem less strong. Look away at something neutral, and it will appear to have the opposite sign. For motion, this means you adapt to one motion for a long time, and when you look away you will see stationary things appear to move. The existence of specialized motion sensitive units is dramatically illustrated by the spiral motion aftereffect. Steady objects will appear to contract after watching an expanding spiral. Adaptation to the moving stimulus causes an aftereffect of perceived motion where there is actually only stationary images. A rotating spiral provides a strong stimulus for adapting motion detectors. Stare at the center of a rotating spiral for about one minute to fatigue motion mechanisms that respond to the expanding pattern. Then shift gaze to some stationary object and watch it appear to contract without getting smaller! See the demo DriftingSpiral.avi Motion Can Break Camouflage Our sensitivity to motion can allow us to see things that otherwise would blend in with the surroundings. Many creatures have colors and patterns that make them blend in with their background, but when they move our visual system is able to see the shape clearly. In the laboratory, we can make stimuli that are perfectly camouflaged except for motion by using completely random colors in the object and the background. These stimuli are called Random Dot Kinematograms. An example is in the demo movie RandomDotUHCO.avi Global Processing in Motion Perception Our perception of motion in the world is built up from individual detectors at each location in the visual field. A single local detector may often have ambiguous motion information, but higher-level mechanisms are usually able to combine all the local motion signals into a unified percept. There are long range interactions across the visual field that “bind” local motions together into a single large object moving in one direction. See the demos DiaMovieG.mov and DiaMovieW.mov for an example of local and global motion perception being very different. In the demo movies, the diamond moves left and right behind some occluding bars. If the bars are visible, the brain connects the four parts of the diamond and sees one object. If the bars are less obvious, the brain sees four independent lines moving in different directions. Motion perception requires the brain to put together local motion signals into one sensible global motion pattern, and sometimes this is ambiguous. The “Aperture Problem”: The individual parts of the moving diamond are seen by neurons in the cerebral cortex as isolated line segments moving through the cell’s “receptive field”: that part of the visual field that activates the particular cell. Seen through this small aperture, the motion always appears to be going perpendicular to the line. Only by combing the activity of many such cells can the brain extract the whole pattern motion. The white circle represents the receptive field of a hypothetical neuron in primary visual cortex. This neuron is not sensitive to areas outside the circle (in gray) but only that part of the visual field that falls inside the white circle. When a straight line or edge moves through the visual field, the cell sees the direction of motion as perpendicular to the orientation of the line, regardless of the true direction of the pattern. On the left, the line moved vertically, on the right it moved horizontally, but the one cell cannot tell the difference. Motion processing in cortex: Area V1 (also called “Area 17”) in cortex is the primary visual area, and has cells with motion selectivity, but they cannot solve the aperture problem. The Middle Temporal area (“area MT”) in the Temporal lobe collects the responses of all those V1 cells and has cells that respond to the whole pattern, rather than small pieces. Experiments suggest that this area has the closest association to our perception of motion in complex displays. Damage to this part of the brain leads to a condition of “motion blindness” or Akinetopsia. Patients with this condition will see objects clearly, and be able to indicate position, but are unable to smoothly follow moving objects or judge their speed. A nearby, related area called the Middle Superior Temporal area (MST) has similar sensitivity to global motion, but also includes information about the rotation of the eyes to help us track moving objects. Motion on the retina produces a flickering signal in individual cones (grating through holes movie). Motion cannot be detected if the local flicker rate is too high or if contrast is too low. Neurons in the cortex compare flicker signals to find the local motion. Local motion is ambiguous (shifting diamond movie). Local motion is adaptable (spiral aftereffect movie). Neurons in specialized regions of the cortex compare local motion signals to find the global motion. Global motion lets us identify objects from their motions (UHCO in random dots). Damage to these brain areas can make you motion blind.